The genus Fuscidea Fuscideaceae, lichenized · PDF fileThe genus Fuscidea (Fuscideaceae,...

The Lichenologist 40(4): 295–328 (2008) � 2008 British Lichen Societydoi:10.1017/S0024282908007755 Printed in the United Kingdom

The genus Fuscidea (Fuscideaceae, lichenized Ascomycota) inNorth America

Alan M. FRYDAY

Abstract: The species of the genus Fuscidea occurring in North America are revised. Two newspecies, Fuscidea appalachensis Fryday and F. texana Fryday, are described from eastern NorthAmerica and southern Texas, respectively. Three new combinations are also made in Fuscidea:Lecidea aleutica Degel. is shown to be a distinct species and not a synonym of Fuscidea lowensis (H.Magn.) R. Anderson & Hertel as previously suggested and is here recognized as F. aleutica (Degel.)Fryday; non-sorediate, apotheciate specimens from eastern North America previously referred to F.recensa (Stirt.) Hertel, V. Wirth & Vezda are recognized as Fuscidea recensa var. arcuatula (Arnold)Fryday; and Fuscidea scrupulosa (Eckf.) Fryday is shown to be the correct name for Fuscidea subreagens(H. Magn.) Oberholl. & V. Wirth. Fuscidea subfilamentosa (Zahlbr.) Brako is shown to be a memberof the Lecidea hypnorum group and the new combination Lecidea subfilamentosa (Zahlbr.) Fryday ismade, and Lecidea gyrodes H. Magn., described from Tennessee, is shown to be a synonym of F.recensa var. arcuatula. Fuscidea cyathoides (Ach.) V. Wirth & Vezda, F. kochiana (Hepp) V. Wirth &Vezda and F. lightfootii (Sm.) Coppins & P. James are considered not to have been correctly reportedfrom North America.

Key words: Appalachian-Great Lakes distribution, Lecidea hypnorum group, lichens, newcombinations, new species

Introduction

The genus Fuscidea V. Wirth & Vezda(1972) was erected for the well-definedLecidea rivulosa group, which had previouslybeen monographed by Magnusson (1925).The genus is characterized by havingTeloschistes-type asci, simple (rarely 1–septate), hyaline ascospores that sometimesbecome brown on maturity, and havingonly brown (fuscous) apothecial pigments.The identity of this pigment is unclear. Itagrees with Leptoclinoides-brown (Meyer &Printzen 2000) in that it becomes dull brownwith a slight olivaceous tint in 10% KOH,whereas it is unchanged in 10% HCl, butbecomes orange-brown in 50% HNO3.However, the pigment does not dissolve inKOH, although a brown solution is some-times obtained suggesting that the pigment

is at least partially dissolved. Fuscidea isfurther characterized by having a photobiontthat divides by binary fission (B. J. Coppinspers. comm.), resulting in a distinctivearrangement of cells in groups of 2, 4 or 8(Kantvilas & McCarthy 2003; fig. 3F).Although this type of photobiont alsooccurs in some other genera of lichenizedfungi, especially in the Trapeliaceae Hertel(Lumbsch et al. 2007; e.g., Trapelia M.Choisy and Trapeliopsis Hertel & Gotth.Schneid.) it is a useful character for confirm-ing the placement in the genus of specimenswithout apothecia.

Galloway (2007) has suggested thatFinerania C. W. Dodge (Dodge 1970),described from New Zealand, is an earliername for Fuscidea. If so, then a case will haveto be made for the conservation of Fuscidea,which was described after Finerania. How-ever, Fuscidea has been widely used sinceits description, whereas Finerania hasnever subsequently been noted by any

A. M. Fryday: Herbarium, Department of PlantBiology, Michigan State University, East Lansing, MI48824-1312, USA. Email: [email protected]

lichenologist since Dodge’s original descrip-tion. Dodge’s description is also factuallyinaccurate as he refers to the photobiont asbeing Trentepohlia.

Traditionally, Fuscidea has, with some res-ervations, been included in the TeloschistalesD. Hawksw. & O. E. Erikss., but molecularwork has suggested that its true affinity lieselsewhere (Lutzoni et al. 2004; Reeb et al.2004; Wedin et al. 2005), and recent work(Bylin et al. 2007; Miadlikowska et al. 2006)has shown that it should be included inthe Umbilicariales J. C. Wei & Q. M. Zhou(Miadlikowska et al. 2006; Zhou & Wei2007), along with other genera of the Fusci-deaceae Hafellner (e.g., Maronea A. Massal.,Orphniospora Korb., and Ropalospora A.Massal.), the Umbilicariaceae Chevall. (i.e.,Umbilicaria Hoffm. and Lasallia Merat), theOphioparmaceae R. W. Rogers & Hafellnersensu Wedin et al. (2005) (i.e., HypocenomyceM. Choisy, Ophioparma Norman and Boreo-placa Timdal) and the Elixiaceae Lumbsch(i.e., Elixia Lumbsch).

The distribution of the genus is interestingin that, although it occurs on all sevencontinents, the non-sorediate species, atleast, often exhibit a high degree of ende-mism, rarely being reported from more thanone region. For example, the commonEuropean species F. cyathoides (Ach.) V.Wirth & Vezda, F. kochiana (Hepp) V.Wirth & Vezda, and F. lygaea (Ach.)V. Wirth & Vezda have not been reliablyreported from North America [although F.cyathoides was reported from Japan by Inoue(1981a)], whereas other European taxahave been reported only from the PacificNorthwest [i.e., F. intercincta (Nyl.) Poelt,and F. mollis (Wahlenb.) V. Wirth & Vezda].Interestingly, this is largely in line withother genera of the Umbilicariales, in whichEuropean taxa are largely absent from borealeastern North America. However, othergenera in the Fuscideaceae (e.g., Ropalospora,Orphniospora) do occur in eastern NorthAmerica and, as North American records ofEuropean species of Umbilicaria are mainlyfrom northern regions, and Fuscidea collec-tions from northern Canada are infrequent,it may be that these species occur there.

Fuscidea has been revised in Japan byInoue (1981a), who also carried out a ‘pre-liminary revision’ of the extra-Japanesespecies (Inoue 1981b), in Europe byOberhollenzer & Wirth (1984, 1985, 1990)and the sorediate, corticolous species byTønsberg (1992), and in Tasmania andAustralia by Kantvilas (2001, 2004). Inaddition, Hertel (1974) and Kalb (1986)transferred species from Colombia andVenezuela, and Brazil respectively, to thegenus, and Breusse (1989) described a newspecies from South Africa. In NorthAmerica, Brako (1991) transferred one cor-ticolous species to the genus (F. subfilamen-tosa (Zahlbr.) Brako; but see below),Tønsberg (1993) reported two recentlydescribed sorediate, corticolous species (F.arboricola Coppins & Tønsberg and F. pusillaTønsberg) from the continent, Brodo &Wirth (1994) revised the species ocurring onthe Queen Charlotte Islands and describeda new species (F. thomsonii Brodo & V.Wirth), and Ekman (1993) provided someuseful anatomical information in his studyof the closely related Ropalospora chlorantha(Tuck.) S. Ekman. Otherwise, the genus hasbeen largely unstudied on the Americancontinent.

In the course of field work on Katahdin,Maine (Dibble et al. unpublished, A. L.Hinds et al. unpublished, Fryday 2006), theauthor made numerous collections referableto the genus Fuscidea. While attempting toidentify and determine the correct names forthese collections, it became evident thatthere were a number of nomenclatural andtaxonomic problems attached to the genusin the region. This paper is an attempt torectify some of these problems.

Materials and Methods

Collections from the following herbaria were studied:ABL, ASU, BG, CANL, DUKE, E, F, FH, H, MICH,MIN, NY, QFA, S, PH, US, UPS, W and WIS alongwith material from the private herbaria of D. Flenniken(Wooster, OH), J. Gagnon (Quebec), E. Lay (Boston),J. Lendemer (Philadelphia), G. Perlmutter (ChapelHill, NC), and G. Thor (Uppsala). Additional distri-bution data was obtained from BG (F. arboricola and F.pusilla) and NY (F. arboricola).

296 THE LICHENOLOGIST Vol. 39

Apothecial characteristics were examined by lightmicroscopy on hand-cut sections mounted in water or10% KOH (K). The ascus structure was studied in0·15% aqueous IKI, both without prior treatment andafter pretreatment with 10% KOH. All ascosporemeasurements were made in 10% KOH. TLC followsthe methods of Culberson & Kristinsson (1970).

Additional comparative material examined: Fuscideacacuminum (Vain.) M. Inoue. Finland: Hogland: onrock, Brenner (H-Nyl—holotype).

Fuscidea circumflexa (Nyl.) V. Wirth & Vezda.Russia: Siberia: Fretum Beringii, Lawrence Bay, 1879,E. Almqvist (H-Nyl—lectotype, S—isolectotype).

Fuscidea fulva (Malme) Kalb.—Brazil: Minas Gerais:Serra do Espinhaco, Sierra do Caraca, osterlich desKlosters Caraca, in einem Cerrado, 1380 m., 8 7 1978,K. Kalb & G. Plöbst (MIN—Lichens Neotropici Exsiccati#369).

Fuscidea impolita (Mull. Arg.) Hertel.—FalklandIslands: East Falklands: Stanley, outcrops along GoatRidge, UTM: 21F VC 2971, 600 ft, 1968, Imshaug(41517) & Harris (MSC).—New Zealand: CampbellIsland: Rock outcrops and feldmark on summit of MtDumas (500 m), 1970, Imshaug 46993 (MSC); ibid.,outcrops above Venus Cove on lower slopes of MtHoney, 1970, Imshaug 47093, 47096 (MSC).

Fuscidea ramboldioides Kantvilas.—Australia:Tasmania: Crandle Mts, Lake St Clair NP, HounslowHeath, summit knoll, 41(18#S, 145(55#E, 1160 m, onlarge rock outcrop in alpine heathland, 2002, G.Kantvilas 554/02 (HO).

Fuscidea subasbolodes Kantvilas.—Chile: Prov.Magallanes: Isla Desolacion, Moorland on hill, S sideof Caleta San Jose, Bahia Tuesday, on hill, 1969,Imshaug (44700, 44722) & Ohlsson (MSC).—FalklandIslands: East Falklands: Stanley, at summit of Mt Kent,cliffs on rock dome, 1968, Imshaug (40438, 40467,40477) & Harris (MSC).—New Zealand: AucklandIsland: summit of Cloudy Peak, 1973, Imshaug 57561(MSC).

Fuscidea cf. umbricolor (Nyl.) Hertel.—Brazil: MinasGerais: Catas Altas, Serra do Caraca, Parque Natural doCaraca, near monastery Santuario do Caraca, 20(06#S,43(29#W, 1300 m, on sandstone, 1997, A. Aptroot40710 (MSC).

Fuscidea sp. Dominican Republic: Summit of PicoTrujillo [Pico Duarte], Cordillera Central (MacisoCentral, Sierra de la Pelonia), 10 000 ft, pinewoodland, 1958, Imshaug 23566 (MSC).

Taxonomy

Fuscidea V. Wirth & Vezda

Mycobank: MB2022.

Beitr. Naturk. Forsch. Südwestdeutschl. 31: 91 (1972);type species: Lecanora austera Nyl., Flora 57:309 (1874)=Fuscidea austera (Nyl.) P. James inHawksworth, James & Coppins, Lichenologist 12: 106(1980).—Lecidea contigua var. rivulosa f. aggregata Flot.,

Flora 11: 674 (1882)=Lecidea aggregata (Flot.) H.Magn., Göteborgs Kgl. Vetensk.-och Vetterh.-Handl., ser4, 29: 12 & 20 (1925)=Fuscidea aggregata (Flot.) V.Wirth & Vezda, Beitr. Naturk. Forsch. Südwestdeutschl.31: 92 (1972).

?Finerania C. W. Dodge, Nova Hedwigia 19: 459(1971); type species: Finerania neozelandica C. W.Dodge, Nova Hedwigia 19: 459 (1971).

Thallus crustose, grey to brown or green-ish, cracked-areolate or areolate, prothallususually present; medulla I� or I+ violet.Photobiont chlorococcoid, Chlorella-type,cells dividing by binary fission.

Apothecia lecideine, usually sessile or ad-nate, black or brown, usually with persistentproper margin, 0·1 mm wide in youngapothecia 0·05 mm in older ones, thallinemargin absent (except in intercincta group).Hymenium colourless, I�; hamathecium ofsimple or sparingly branched paraphyses,1·5–2(–3) �m wide, septate, often monili-form above, apices scarcely enlarged orswollen to 5 �m with a brown cap, lax inK; epihymenium brown, 5–20 �m tall.Hypothecium colourless. Exciple of radiatinghyphae, cortical cells with brown pigment,usually paler within but occasionally uni-formly brown. Asci 8-spored, Teloschistes-type; ascospores colourless but oftenbecoming brown with age, usually simplebut occasionally with a median constrictionor septum (1-septate).

Conidiomata pycnidia, often present,immersed in thalline warts or becomingemergent; conidia cylindrical or bacilliform.

Chemistry. Divaricatic acid in manyspecies. Also alectorialic, fumarprotoce-traric, norstictic and sekikaic acids in somespecies.

Fuscidea aleutica (Degel.) Frydaycomb. nov

Mycobank: MB 511668.

Lecidea aleutica Degel., in Meddel. Göteborgs Bot.Trädgard 12: 111 (1938); type: USA, Alaska, AleutianIslands, Unalaska Island, 9 v 1932, E. Hultén 5319(UPS—holotype (L74992) 132889!).

(Figs 1A, 2 & 3)

2008 Fuscidea in North America—Fryday 297

Thallus brown, thin, cracked-areolate,with a thin black prothallus between adja-cent thalli; medulla I+ violet.

Apothecia black, �adnate, not constrictedbelow, orbicular, 0·3–0·4(–0·5) mm diam.;disk black, matt, flat; proper margin persist-

ent, slightly raised, 0·05–0·1 mm thick.Thecium 70–80 �m tall, paraphyses simple,upper 20–25 �m brown pigmented, apicalcells slightly swollen (3 �m); excipulumannular, intercincta-type (see below) poorlydeveloped, cortex brown, paler within but

F. 1. Thallus and apothecia or soralia of Fuscidea species. A, F. aleutica (holotype); B, F gothoburgensis (Dibble11001); C, F. lowensis (Fryday 8610); D, F. mollis (Räsänen s.n.: Lich. Fenn. Exs. 974); E, F. thomsonii (Brodo

17833); F, F. kochiana (Fryday 6073). Scales: A, E & F =0·5 mm; B, C & D=1·0 mm.


obscured by brownish granules that dissolvein K. Asci 75–85�12–15 �m, cylindrical toslightly clavate; ascospores simple, 8–10�6–7 �m, subglobose to broadly ellipsoid.

Conidiomata not observed.

Chemistry. K�, C�, Pd�, UV+ white(medulla). Divaricatic acid by TLC.

Distribution and Ecology. Fuscidea aleuticais known from only the type collection fromUnalaska Island in the Aleutian Islands,Alaska, and from three collections from theQueen Charlotte Islands, British Columbia,but is probably more widely distributedalong the NW coast of North America. Thethree collections from the Queen CharlotteIslands were mentioned by Brodo & Wirth(1998) as possibly an unnamed species or asaberrant forms of F. thomsonii.

Remarks. This species is characterized byhaving a thallus containing divaricatic acidand an amyloid (I+ violet) medulla, andapothecia with only a lateral (annular)intercincta-type exciple (Oberhollenzer &Wirth 1984). In the intercincta-type exciple,

the proper exciple is poorly developed ornon-existent (in immersed apothecia)whereas a pseudothalline exciple is devel-oped from vertical thalline hyphae, theupper 20–30 �m of which are dark-brownpigmented closest to the apothecium givingthe appearance of a thick, dark properexciple surrounded by a thin thalline margin.The inner edge of this pseudothalline marginoften has a white pruina, giving the appear-ance of a ‘‘halo’’ around the apothecial disc(Fig. 3).

Fuscidea aleutica is very similar to F. thom-sonii, with which it is sympatric, and F.impolita from the subantarctic. All threespecies have a thin thallus with an amyloidmedulla, and �adnate apothecia with only alateral exciple. Fuscidea impolita is anatomi-cally distinct in having more robust paraphy-ses with a darker pigmented cap and withmuch shorter asci (25–30 �m), but the onlyanatomical difference between F. aleuticaand F. thomsonii appears to be that F. aleu-tica has more sessile apothecia with a betterdeveloped exciple as well as a more intenselypigmented exciple and paraphyses. It ispossible that these differences, and those ofthe gross morphology, are environmentallyinduced, and only one taxon is involved, inwhich case F. aleutica has nomenclaturalprecedence. However, when the two speciesoccur together (e.g., Brodo 17833; Fig. 3)they remain distinct, and as I have not seeneither taxon in the field I prefer, at this stage,to maintain them as distinct species.

When making the new combination Fus-cidea impolita, Hertel (1984) mentioned thatLecidea aleutica Degel. appeared to be closelyrelated, if not identical, to F. lowensis (H.Magn.) R. Anderson & Hertel and thisopinion has been followed in subsequentNorth American lichen checklists (Egan1987; Esslinger 1995; Esslinger 1997).Hertel was probably of the opinion that thetwo species were closely related becauseboth species (along with F. incompta andF. umbricolor (Nyl.) Hertel from Colombiaand Venezuela) have an amyloid medulla,whereas most other species of the genusdo not (e.g., Oberhollenzer & Wirth 1984;Kantvilas 2001). However, two further

F. 2. Known distribution of Fuscidea aleutica (●) andF. thomsonii (�), and North American distribution ofF. mollis (:) and F. intercincta. Shaded area (QueenCharlotte Islands), all three species plus F. intercincta.


species with an amyloid medulla have nowbeen described (F. submollis Inoue, and F.thomsonii), and it is clear that this characteris not as critical as Hertel believed. In fact, F.aleutica differs from F. lowensis in a numberof characters. It has a thin, cracked areolate-rimose thallus and flat,�adnate apotheciawith a thick persistent margin (Fig. 1A),contrasting with the thick areolate thallusand convex immarginate apothecia of F.lowensis (Fig. 1C). Internally, the excipulumof F. aleutica does not extend under thehypothecium, whereas that of F. lowensisdoes, and the paraphyses are also far lessconglutinate than those of F. lowensis, and atleast some are branched and anastomosing,unlike those of F. lowensis, which are alwayssimple. The two species are also allopatricand ecologically distinct: F. aleutica is con-fined to the coast of the Pacific NW (Fig. 2),

whereas F. lowensis occurs only at higheraltitudes in NE North America (Fig. 9).

Additional specimens examined. Canada: BritishColumbia: Queen Charlotte Islands, Graham Island,Port Channal off Athlow Bay, on south side of inlet(Goose Cove) shoreline rocks and trees and bluffs, onboulder at summit, 320 ft, 1967, I. Brodo (10444 p.p.),M. Shchepanek & W. Schofield (CANL); ibid., in aero-haline, 1967, I. Brodo (10455), M. Shchepanek & W.Schofield (CANL); ibid., head of Bigsby Inlet, 52(38#N131(46#W, along cascade and in forest along south sideof first lake west of inlet, Tsuga-Thuja forest, on rocks(top surface) at edge of stream, 1971, I. Brodo (17833),P. Wong & W. Schofield (CANL).

Fuscidea appalachensis Fryday sp.nov.


Fuscideae kochianae similis sed areolis bullatis et sorediainstructis, apotheciis non immersis prominentermarginatis.

F. 3. Fuscidea aleutica (left) and F. thomsonii (right) growing together on the Queen Charlotte Islands (Brodo17833). Scale=1 mm.


Typus: USA, Maine, Piscataquis Co., Baxter StatePark, Katahdin, top of Tabor Gully, 45(55.59#N,68(55.73#W, 1415 m, damp rocks on ledge at side ofgully, 4 September 2004, A. M. Fryday 8897(MSC—holotype).

(Figs 4–7)

Thallus (Fig. 4) effuse rarely more than1–2 cm diam. but with adjacent, confluentthalli sometimes covering a larger area,0·15–0·2 mm thick; pale grey, composed offlat to strongly convex, �contiguous areoles(2–)4–6 mm across; cortex 20–25 �m thick,hyaline with minute brown granules; soralia(Fig. 5) infrequent in fertile areas of thallusbut often abundant elsewhere, �discrete,c. 0·1 mm diam., but often coalescing toform aggregate soralia 0·3–0·5 mm diam.;soredia farinose, 0·01–0·02 mm diam., sur-face soralia pale brown, green internally;medulla I�. Photobiont Chlorella-type, cells8–12(–15 when dividing) �m diam.; dividingby binary fission.

Apothecia (Fig. 5) frequent or lacking,�immersed to (usually) adnate, not con-stricted below, orbicular when youngbecoming flexuose or occasionally linearwhen mature, 0·7–0·9 mm diam.; disc paleto dark brown (rarely black), often withdense grey pruina; proper margin persistent,slightly raised, 0·1 mm wide, concolorouswith the disc or paler, not pruinose but withwhite margin on inner edge when young,

often deeply radially striate, especially inyoung apothecia. Thecium 50–60 �m tall;epihymenial layer brown, 25–30 �m tall, withminute brown granules. Paraphyses simple,lax in K, septate, 2–2·5 �m wide, notwidening at the apex but with a dark browncap. Hypothecium hyaline, 150–200 �m tall,composed of randomly arranged hyphae notclearly separated from the thecium. AsciTeloschistes–type, cylindrical, 45–50�8–10(–12) �m; ascospores hyaline, becomingbrownish when over mature, globose tosub-globose, 8–9�6–7 �m, 8/ascus, �uni-seriate in ascus. Excipulum cupular, c. 25–30 �m thick; composed of radiating hyphae5 �m wide, inner part hyaline, outer 15–20 �m with minute brown granules, corticalcells with dark brown pigmented walls.

Conidiomata pycnidia (Fig. 6), frequent,immersed in the thallus, 0·05–0·1 mmdiam., pale brown, with a dark brown‘involucrellum’ developing in older pycnidia,0·2–0·25 mm across; conidia ellipsoid 4·5–5·0�1–1·5 �m.

Chemistry. K�, C�, KC�, Pd�, UV+white. Divaricatic acid by TLC.

Distribution and ecology. Fuscidea appa-lachensis appears to be restricted to theAppalachian Mountains of eastern NorthAmerica, where it has been collected from

F. 4. Fuscidea appalachensis at 1400 m on Katahdin, Maine (photo A. Dibble).


New Brunswick in the north to Tennessee inthe south (Fig. 7). It appears to be particu-larly frequent in the Adirondack Range ofNew York, but that may be because ofgreater collecting activity in that area.

Fuscidea appalachensis is a species ofexposed, sunny, hard, siliceous rocks(Fig. 4), usually above the tree-line, whereit often grows with other species of thegenus.

F. 5. Fuscidea appalachensis. A, apothecia (holotype); B, apothecia and soralia (holotype); C, soralia (Fryday8844); D, soralia (Fryday 8873). Scales: A–D=0·5 mm.

F. 6. Fuscidea appalachensis; Pycnidia (holotype). Scales: A & B=0·1 mm.


Remarks. In the past, F. appalachensis hasusually been identified in North America asFuscidea kochiana (e.g., Lowe 1939, andprobably Eckfeldt 1895) but that species hasimmersed, immarginate apothecia (Fig. 1F)and has not been correctly reported fromNorth America. In the absence of soralia,F. appalachensis could be confused with F.mollis (Wahlenb.) V. Wirth & Vezda, butthat species has sessile apothecia (Fig. 1D)and a thallus composed of flat areoles. Fus-cidea appalachensis could also be mistaken forF. recensa var. arcuatula, but that species hasvery different ascospores (Fig. 12), and alsoa more heavily pigmented exciple and para-physes with the top two cells pigmented, notjust a pigmented cap as in F. appalachensis.In the absence of apothecia, F. appalachensisand F. recensa var. recensa are difficult toseparate, although if pycnidia are presentthey can be separated by F. appalachensishaving immersed pycnidia whereas those ofF. recensa var. recensa are raised. Also, F.appalachensis tends to have a thicker thallus,is not known to occur at low altitudes andappears to be confined to eastern NorthAmerica, whereas F. recensa var. recensa ismost frequent in central USA.

A further collection from Katahdin(Fryday 8870) is also possibly referable tothe new species. It differs in having moresessile, strongly gyrose/contorted apotheciabut is otherwise similar. Also from the samelocation is a collection with sessile apotheciathat resembles F. cacuminum (Vain.) M.Inoue, but the holotype, and only collectionof that species is very poor and morematerial is required to determine whetherthe two collections are conspecific.

Additional specimens examined. Canada: NewBrunswick: Northumberland Co., Mount CarletonP.C., summit of Mount Carleton, 47(24#N 66(53#W,820 m, on rock, 11 Aug 1988, E. Haber & H. R. Hindss.n. p.m.p. (CANL).—USA: Maine: Piscataquis Co.,Baxter State Park, Katahdin, North Basin, 45(55.76#N68(54.86#W, 975 m, granitic boulder, 2004, Fryday8844 (MSC); ibid., just east of the ‘calcareous seep’,45(55.78#N 68(55.50#W, 1120 m, on exposed, verti-cal rock face with no calcareous influence, 2004, Fryday8866, 8872, 8873 (MSC). New Hampshire: Coos Co.,White Mountain NF, Mt Washington, along AutoRoad at Cragway Spring, along Nelson Crag Trail,44(17#N 71(18#W, 1430–1465 m, heath and krumm-holz vegetation on leeward slope, 1999, R. Harris 4319p.p. (NY). New York: Essex Co., Mount Marcy (nearLake Placid), 5300 ft, on rocks on N side, 1933, J. Lowe2604 (MICH); ibid., on exposed rock, 1933, J. Lowe2888 p.m.p. (FH); ibid., 5300 ft, on rocks, 1933, J.Lowe 2946 p.p. (MICH); ibid., 5300 ft, on rocks northexposure, 1933, J. Lowe 3037 (MICH); ibid., 4900 ft,on rocks in gully, 1933, J. Lowe 3187 (MICH); ibid.,summit 5110 ft in alpine zone, at summit 5100 ft, 1963,I. Brodo 4768 (MSC); ibid., Adirondack Mts, Algon-quin Peak, McIntyre Mt, south of Lake Placid, 5100 ft,on summit above tree line, 1963, C. Wetmore 12288p.p. (MIN); ibid., lower part of alpine zone, on rocks,1963, J. Thomson 11234 (WIS). North Carolina: BurkeCo., Linville Gorge Wilderness Area, on exposed cliffface, 1997, P. Smith 3114 (NY). Tennessee: GreatSmoky Mountains, near Alum Cave, c. 1500 m. 13 ix1939, G. Degelius s.n. p.p. (US). Virginia: Giles Co.,Bald Knob near Mountain Lake, on exposed rocks,22 iii 1975, D. Fahselt s.n. (DUKE). West Virginia:Tucker Co., Monongahela National Forest, Dolly SodsWilderness, Bear Rocks, 39(03#58$N 79(18#06$W,1205 m, ericaceous heath and sandstone exposures,2001, W. Buck 38985 (NY).

Fuscidea arboricola Coppins &Tønsberg

Mycobank: MB358665.

In Tønsberg, Sommerfeltia 14: 134 (1992); type: Norway,Nordland, Bindal, Skjelsvik, UTM 33W UN 7131(1825 III), 0–10 m., on trunk of Betula pubescens, 1 June

F. 7. Known distribution of Fuscidea appalachensis.


1982, Tønsberg 6865 (BG—holotype; E, UPS—isotypes).

(Fig. 8)

Thallus areolate to warted, green-brown;soralia pale yellow-green, initially discrete,becoming confluent towards the centre;distinct, brown prothallus visible betweenareoles and at margin; medulla I�.

Apothecia rare, sessile, disc dark brown,proper margin often becoming flexuose.Ascospores simple or 1-septate, with a medianconstriction, 7–9�4–5 �m.

Chemistry. K�, C�, Pd+ red, UV�.Fumarprotocetraric acid by TLC.

Distribution and ecology. Fuscidea arboricolais a frequent, corticolous species in easternNorth America, where it has anAppalachian-Great Lakes distribution.Fertile material is known from Maine,Massachusetts and Newfoundland.

Remarks. A usually sterile, sorediate corti-colous species containing fumarprotoce-traric acid (Pd+ red). It was first reportedfor North America by Tønsberg (1993)from Newfoundland, New York and NorthCarolina, although numerous earlier collec-tions are now known (mostly misidentifiedas F. lightfootii (Sm.) Coppins & P. James),including two fertile specimens collected byHenry Willey from Massachusetts in 1868and 1869 (in US).

Fuscidea arboricola is the only NorthAmerican species of the genus containingfumarprotocetraric acid (F. cyathoides, ausually saxicolous species, also containsfumarprotocetraric acid but is considerednot to occur in North America; see below).

Selected specimens examined. Canada: Newfoundland:Newfoundland Island, Avalon Peninsula, ButterpotProvincial Park, east side of Big Otter Pond, 47.399(N53.043(W, c. 160 m., mesic to wet coniferous forest,2007, J. Lendemer 10153 (NY: fertile). Ontario: AlgomaDistrict, Lake Superior P.P., Crescent Lake camp-ground, 47(17#N 84(33#W, maple forest, on Betulapapyrifera, 1993, S. D. & S. Sharnoff 1106.16 (CANL);Bruce Co., 2 miles SE of Miller Road, on Hwy 6,45(04#N 81(24#W, Abies-Thuja forest, on Abies balsa-mea, 1976, P. Wong (2039) & P. Bowler (CANL);Renfrew Co., SW of Kennellys Mountain, 45(17#N76(54#W, deciduous forest, on Ostrya virginia, 1985, P.Wong 3777 (CANL); Lennox & Addington Co.,Centerville, 10 miles NE of Napanee, Basswood Hill,44(25#N 76(55#W, 150 m, on sugar maple in decidu-ous forest, 1977, P. Wong 2494 (CANL); Simcoe Co.,Hillsdale, SE of town, 44(35#N 79(45#W, deciduouswoods, on young tree, 1984, P. Wong (3526, 3530,3532) & J. Krug (CANL). Quebec: Gatineau, Aylmer,N of Pink Road, E of Vanier Road, 45(25#N 75(23#W,mature lowland Thuja occidentalis stand, on Acerrubrum, 1995, I. Brodo (28912) & F. Brodo (CANL).—USA: Maine: York Co., Alfred, Massabesic Experi-mental Forest, Chamaecyparis swamp, 43(26#53$70(40#18$W, 60 m, on branches of Chamaecyparis,2002, R. Harris 46244 (hb. Lendemer, ex NY—fertile).Massachusetts: [Bristol Co.,] New Bedford, 1868, H.Willey (US—fertile); ibid., Weymouth, 1869, H. Willey(FH, US—fertile); Norfolk Co., Blue Hills Reservation,Canton, 25 vii 2000, E. Kneiper, S. LaGreca & D.Greene (FH). Michigan: Kalkaska Co., Kalkaska,Kalkaska RV Park & Campground, 44(42.6#N,85(10.0#W, 330 m, Acer, 2007, Fryday 9130 (MSC).Minnesota: Lake Co., Boundary Water Canoe AreaWilderness, North of Gabbro Lake, 47(52#28$N,91(35#25$W, mixed upland forest, 2003, P. Johansson51, 52, (MIN). West Virginia: Tucker Co.,Momongahele National Forest, Olsen’s Bog, 39(07#N,79(36#W, 1000 m, at edge of bog, on Rhododendronbranches, 2001, S. LaGreca 795 (FH).

F. 8. Known North American distribution of Fusci-dea arboricola (●), and F. gothoburgensis (:). Includesone record of F. arboricola from North Carolina notseen by the author that was collected and identified byT. Tønsberg (BG), and several from northeast NorthAmerica from New York Botanic Garden’s ‘‘VirtualHerbarium’’ <http://sciweb.nybg.org/science2/Virtual

Herbarium.asp>, also not seen by the author.


Fuscidea gothoburgensis (H. Magn.)V. Wirth & Vezda

Mycobank: MB342003.

Beitr. Naturk. Forsch. Südwestdeutschl. 31: 92 (1972).—Lecidea gothoburgensis H. Magn., Kongl. GötheborgskaWetensk. Samhällets Handl., Wetensk. Afd. 29: 15(1925); type: Sweden, Goteborg, Botanical Garden, onsteep rocks, 1925, A. H. Magnusson 7614a (UPS—lectotype, fide Oberhollenzer & Wirth 1985: 2).

(Figs 1B & 8)

Thallus pale grey, areolate; areoles eitherdispersed or contiguous, flat to slightly con-vex; soralia usually concolorous with thethallus, rounded, often delimited by a raisedmargin; prothallus black, well-developedbetween areoles and at margin; medulla I�(rarely I+ violet).

Apothecia uncommon; unknown in NorthAmerican collections; generally small, 0·3–0·9 mm diam., but occasionally larger,sessile. Ascospores broad ellipsoid, 7–10�5–7 �m.


Distribution and ecology. Fuscidea gotho-burgensis is a saxicolous species that wasrecently reported for North America from asingle collection from Katahdin, Maine(Fryday 2006), but earlier, unidentified, col-lections have since been discovered fromQuebec (see below).

Remarks. Fuscidea gothoburgensis is charac-terized by a sorediate thallus composed of�confluent areoles containing divaricaticacid, and with small apothecia, althoughthese are not known in North Americanmaterial. Forms consisting of dispersedareoles on a black hypothallus have beenrecognized as f. maculosa (H. Magn.) Poelt)but, as in the Katahdin collection, thisintergrades with the typical form.

As with other saxicolous species of thegenus it occurs on hard, siliceous rocks, butusually in shaded habitats.

Specimens examined. Canada: Quebec: N side of LakeLouis, c. 80 mi N of Schefferville, on rock, 1990, D.Wehr 39 (with F. scrupulosa) (NY); Monte Otish,Monte Centlans, 22 vii 2003, J. Gagnon (hb. Gagnon);ibid., Monte Maria-Victoria, 24 vii 2003, J. Gagnon(hb. Gagnon).—Great Britain: Scotland: V.C. 88Mid Perthshire: Ben Lawers, Allt a’ Mhoirneas, 27/6137, 250 m, wooded ravine, 1989, Fryday s.n. (hb.Fryday); V.C. 92 South Aberdeen: Braemar, EasternCairngorms SSSI, Mar Lodge Estate, Clais Fhearnaig,37(NO)/066.930, 505 m, acid rock boulder field abovelochan on S side of narrow valley, 2005, Fryday 8979,8985 (MSC). V.C. 105 West Ross-shire: Beinn EigheNNR, Meall na h-Airgh-ardain, 18/9865, 300 m,shaded north-east facing rock face, 1991, Fryday 2768(hb. Fryday).—Sweden: Värmland: Munkfors: Bryn-gelstorp, on vertical rock face, 21 v 1951, S. Sundell s.n.(MSC).—USA: Maine: Piscataquis Co., Baxter StatePark, Katahdin, North Basin, calcareous seep,45(55.78#N, 68(55.50#W, 1120 m, shaded graniticrock face, 2003, A. Dibble 11001 (MAINE).

Fuscidea intercincta (Nyl.) Poelt

Mycobank: MB342005.

Norw. J. Bot. 25: 127 (1978).—Lecanora intercinctaNyl., Flora 64: 531, (1881); type: Portugal, Beira Alta,Serra de Estrella, 8 1881, J. Henriques(H-Nyl—holotype).

(Fig. 2)

Thallus grey, areolate; areoles continuousor dispersed (North American material) on ablack prothallus; medulla I�.

Apothecia immersed or slightly emergent,0·3–0·6 mm diam., thin, pseudothallinemargin present, forming a pale ‘halo’ aroundthick black zone, intercincta-type (cf. F. aleu-tica), disc black, flat, matt. Ascospores broad-ellipsoid, 10–13�7–8 �m.


Distribution and ecology. Fuscidea intercinctais a saxicolous species that was reportedfrom the Queen Charlotte Islands by Brodo& Wirth (1998), although these collectionsdiffer from European ones in having a lesswell-developed thallus and somewhat largerascospores.

Remarks. This species is characterized by agrey thallus containing divaricatic acid and


having a non-amyloid medulla (I�), and bysemi-immersed apothecia with a lateralexciple that have a white ring around theoutside of the disc. The Canadian collec-tions resemble F. thomsonii, but differ inhaving an I� medulla. However, this reac-tion can be variable in other species ofFuscidea (e.g., F. gothoburgensis) and it isnot impossible that these specimens are anaberrant form of F. thomsonii.

Specimens examined. Canada: British Columbia:Queen Charlotte Islands, Moresby Island, TakakiaLake, east side 52(56#N 132(03#W, 1930 ft, alongshore and in fens just back from shore, on verticalface of shoreline rock, 1967, I. Brodo (10920) & M.Shchepanek (CANL); ibid., on boulder at water’s edge,1967, I. Brodo (10909) & M. Shchepanek (CANL).—Great Britain: Scotland: V.C. 98 Argyll Main: GlenCoe, Aonach Eagach, Creag nan Gabhar, G.R. 27/1357, 600 m, top of siliceous boulder, 1992, Fryday3329 (hb. Fryday).—Norway: Møre og Romsdal:Bogeskarnakken, Stadt, 1904, J. Havaas (DUKE);ibid., Dalsbø, Stadt, 17 viii 1911, J. Havaas (DUKE).Nordland: Sandnessjon, 30 m, on a steep rock facing thenorth, 5 vii 1924, A. Magnusson (DUKE: Magnusson,Lichenes selecti scandinavici exsiccati).

Fuscidea lowensis (H. Magn.) R.Anderson & Hertel

Mycobank: MB115835.

In Hertel, Herzogia 5: 450 (1981).—Lecidea lowensis H.Magn., Acta Horti Gothob. 10: 11 (1936); type: USA,New York, Essex Co., Mt Marcy (near Lake Placid),5300 ft, 16 Aug. 1933, J. L. Lowe 2860 (UPS—holotype; US, FH—isotypes!).

(Figs 1C & 9)

Thallus brown, often with a grey pruina,thick, of convex, warted areoles, occasionallythinner with flatter areoles; black prothallusvisible at margin; medulla I+ violet.

Apothecia black, flat, sessile, constrictedbelow 0·4–0·7(–0·9) mm diam.; proper mar-gin persistent, becoming flexuose, 0·05–0·1 mm wide. Thecium 60–70 �m; paraphysessimple, branched above, 1·5–2 �m thickwith apical cell swollen (to 5 �m), upper10–20 �m brown pigmented. Asci cylindrical20–50�12–15 �m; ascospores subglobose,7–10�6–7 �m. Exciple cupular, uniformlydark brown.

Conidiomata: pycnidia, immersed in thal-lus but becoming emergent, black, to0·02 mm diam. with gaping ostiole; conidiabroad ellipsoid 4�2 �m.


Distribution and ecology. Fuscidea lowensis isa saxicolous species widespread in NE NorthAmerica on acidic rocks at higher elevations,reaching almost sea-level near the AtlanticCoast (Newfoundland).

Remarks. This species is characterized by athick dark brown verrucose thallus contain-ing divaricatic acid (by TLC) and with anamyloid (I+ violet) medulla. It wasdescribed by Magnusson (1935) from acollection by J. Lowe from Mt Marcy.

One collection (Harris 53860) fromNewfoundland agrees with F. lowensis ingross morphology and anatomy but has athallus with a non-amyloid medulla (I�)and lacking divaricatic acid. As there areseveral typical collections of F. lowensis from

F. 9. Known distribution of Fuscidea lowensis.


the same locality, I consider it to be ananomolous form of that species.

Selected specimens examined. Canada: New Brunswick:Northumberland Co., Mount Carleton P.C., summit ofMount Carleton, 47(24#N 66(53#W, 820 m, onrock, 11 viii 1988, E. Haber & H. Hinds s.n.(CANL). Newfoundland: Newfoundland Island, AvalonPeninsula, Hawke Hills Ecological Reserve, 47.316(N53.128(W 100 m., on pebble in rocky ground in alpinetundra, 2007, J. Guccion 1187 (MSC), J. Lendemer10057, 10059 (NY), R. Harris 53841, 53860 (NY).Quebec: Parc de la Gaspesie, Mont Albert, pres dusommet nord, 48(55#N 66(10#W, sur roc (amphibo-lite), 1977, M.-C. Hamel 13 (CANL); MistassiniTerritory, Otish Mountains, in subalpine zone on Montdu Lagope [??], 52(18#M 70(24#W, 2800 ft, onexposed rock, 1971, M. J. Shchepanek 71-L-32(CANL); L’Ungava Oriental, Riviere George, LacIndian House, 56(03#N 58(50#–60(00#W, sommet decolline a l’est de la riviere, 1947, J. Rousseau 482(US).—USA: Maine: Piscataquis Co., Baxter StatePark, Katahdin, summit area, 45(54.13#N,68(55.64#W; 1475 m, granitic boulder, 2003, Fryday8610 (MSC); ibid. North Basin, 45(55.67#N,68(54.52#W, 935 m, granite boulder, 2003, Fryday8625 (MSC). New Hampshire: [Grafton Co.,]Franconia Mountains, summit of Mt Moosilauke,4811 ft, 10–17 Aug 1898, C. Cummings (DUKE,MSC, WIS: Decades of North American Lichens Exsiccati#311: as Lecidea tenebrosa); ibid. (FH; LichenesBoreali-Americani #240: as Lecidea tenebrosa); Coos Co.,Presidential Dry River Wilderness Area, knob half mileNE of Mt Crawford along Davis Path, 2950 ft, 1988, C.Wetmore, 61941 (MIN); ibid., on east side of MtMonroe, 5000 ft, 1988, C. Wetmore 62412 (MIN);ibid., White Mountain NF, Mt Washington, along AutoRoad at Cragway Spring, 44(17#N, 71(18#W, 1430–1465 m, heath and krummholz vegetation on leewardslope, 1999, R. Harris 43191, 43193, 43197 (NY). NewYork: Essex Co., summit ridge of Whiteface Mountain,4600 ft, 1983, R. Harris 16612 (NY); Mount Marcy(near Lake Placid), 5300 ft, on face of rock, 1933, J.Lowe 2607, 2665 (MICH); ibid., underside of rock NWexposure, 4900–5300 ft, 1933, J. Lowe 2836 p.p. (FH,syntype of L. kochiana var. subregens); ibid., Lake Placid,Algonquin Peak south of Mt Marcy, 5110 ft [1558 m],1963, I. Brodo 4750 (CANL, MSC), 4768 (MSC).

Fuscidea mollis (Wahlenb.) V. Wirth &Vezda

Mycobank: MB342009.

Beitr. Naturk. Forsch. Südwestdeutschl. 31: 92 (1972).—Lecidea rivulosa � mollis Wahlenb., Fl. Lapp. 472(1812).—Lecidea mollis (Wahlenb.) Nyl., Lich Scand.223 (1861); type: Norway, Finmark, Rypklubb, 8 vi1802, G. Wahlenberg (UPS—lectotype, fide Hertel1997: 197).

(Figs 1D & 2)

Thallus pale grey, regularly and finelycracked, areoles flat, contiguous (groups ofareoles dispersed on a black prothallus insome North American material); black,delimiting prothallus at margin; medulla I�.

Apothecia to 1 mm diam., rounded, dis-crete, sessile. Ascospores broad-ellipsoid,8–10�5–6 �m.


Distribution and ecology. Fuscidea mollis isa saxicolous species correctly reported inNorth America only from the QueenCharlotte Islands and Alaska. The speci-mens from the Queen Charlotte Islandsreported by Brodo (1976) and Brodo &Wirth (1998), have a thinner, more dis-persed thallus, but are otherwise identical toEuropean collections of this species.

Remarks. Fuscidea mollis is characterizedby a smooth, finely cracked areolate thalluscontaining divaricatic acid, and sessileapothecia. In North America, it has beenreported from Labrador (Eckfeldt 1895), theWhite Mountains (Tuckerman 1888), andthe Queen Charlotte Islands (Brodo & Wirth1998). Thomson (1997) also gives localitiesin New York, Nunavut, and Alaska. Ofthe specimens referred to this species inThomson’s herbarium (in WIS), one, fromAlaska, is F. mollis, but the rest are incor-rectly identified. No specimens of F. molliscould be found in PH where Eckfeldt’scollections are housed (J. Lendermer pers.comm.), and other collections from else-where in northeastern North America aremisidentifications of other species, mostly F.appalachensis.

Selected specimens examined. Canada: British Colum-bia: Queen Charlotte Islands, Graham Island offAthlow Bay, 53(35#N 132(55#W, shoreline rocks andbluffs on south side of inlet (Goose Cove), 1967, I.Brodo (10434), M. Shchepanek & W. Schofield (ASU,CANL, DUKE, MSC, WIS—Lichenes CanadensesExsiccati #115); ibid., Moresby Island, ‘‘Blue HeronBay’’, north of Sunday Inlet, bluff above bay, 54(41#N


132(02#W, 500 ft, Pinus contorta stand, on quartz veinon summit boulder, 1968, I. Brodo 14094 (DUKE,CANL); ibid., Pocket Inlet, close to mouth of inlet,52(37#N 131(48#W, in open fens, 1968, I. Brodo(14227), P. Wong & N. Turner (CANL).—Finland:Ostrobottnia borealis: Sino, Kivalot, Alapenikka, adrupen graniticam apricamque, 21 viii 1942, V. Räsänen(MSC—Lichenes Finniae Exsiccati No. 974).—Norway: Finnmark: Fredheim i Sydvaranger [Sør-Varanger, near Fredheim, Alt.: 15–70 m], 1906, J.Havaas (Hav 3319, 3324) (DUKE); Mehavn iFinnmark [Gamvik, Mehavn, Alt.: 1–80 m], 1906, J.Havaas (Hav 3325, 3326) (DUKE); Nordl., Lodingen,vii 1919, E. Vrang (NY).—Sweden: Jämtland: Snasa-hogen, in rupibus apricis, 10 vii 1912, G. Malme(NY—Malme, Lichenes suecici exsiccati #272). LapponicaTornensis, Karesuando, in summo monte Vuokavaara,c. 600 m., ad saxa granitica, G. Lang (NY—Kryptogammae exsiccatae; editae a Museo PalatinoVindobonensi #1955). Torne Lappmark: Jukkasjarvi, inalp. Vassiaive (Vassitjakko), 68(22#0$N, 18(12#0$E, 1Jul. 1927, E. Vrang (ASU).—Korea: Gangwon Prov-ince: Sorak Mts, Sorak-san National Park, Mt Dachong,1·5–2·5 km NNE of Osaeck, 38(05.30–06.45#N,128(27.00–30#E, 700–1400 m, siliceous rocky out-crops, 2006, G. Thor 20264 (KB, UPS).—USA:Alaska: Endicott Mts. of Brooks Range, ArrigetchCreek valley, 67(26#30$N, 154(05#W, 4000 ft, mor-raine in glacial cirque, on granitic rocks, 1980, D.Cooper CL-348 (WIS).

Fuscidea praeruptorum (Du Rietz &H. Magn.) V. Wirth & Vezda

Mycobank: MB342011.

Beitr. Naturk. Forsch. Südwestdeutschl. 31: 92 (1972).—Lecidea praeruptorum Du Rietz & H. Magn., in DuRietz, Zur method. Grundl. d. mod. Pflanzensoz. (Akad.Abh.): 164 (1921); type: Sweden, Bohuslan,Hallesdalen par O} dsmal, on stone fence, 21 vi 1918,A. H. Magnusson (UPS—lectotype, fide Skjolddal 1983:100).

(Fig. 10)

Thallus brown, areolate, delimited by ablack prothallus; soralia initially discrete andpale green to cream-coloured, becomingconfluent and brownish; medulla I�.

Apothecia very rare; sessile with a narrowproper margin, Ascospores 9·5–12�3–4 �m,bean-shaped.

Chemistry. K�, C+ red, KC+ red, Pd+yellow, UV+ orange. Alectorialic acid byTLC.

F. 10. Known North American distribution of Fuscidea praeruptorum. Corticolous (●), saxicolous (�), andknown distribution of F. texana (:).


Distribution and ecology. In Europe, F.praeruptorum is usually a saxicolous speciesof damp, shaded siliceous rocks, althoughseveral corticolous collections were reportedfrom Norway by Tønsberg (1992). In NorthAmerica, however, it is known from severalcorticolous collections from SE Alaska,British Columbia and Newfoundland (inBG), but by only four saxicolous collectionsfrom New York, Newfoundland and WestVirginia. It is probable that this reflects acollecting bias towards corticolous speci-mens rather than a habitat preference for thisspecies in North America.

Remarks. This is the only sorediate speciesof the genus that contains alectorialic acid.Although it usually lacks apothecia, it isfurther characterized by having somewhatcurved, ellipsoid, 9–12�4–5 �m ascospores(Magnusson 1925).

Fuscidea praeruptorum was first reportedin North America from rocks in theAdirondack Mountains of New York byLowe (1939). The specimen (in MICH)now lacks apothecia, although they werepreviously present (Lowe 1939), but has aChlorella-like photobiont and soredia thatare KC+ red, Pd+ yellow, UV+ yellow,indicating the presence of alectorialic acid,and I see no reason to doubt the determi-nation. In addition, I have also seen recentsaxicolous collections from West Virginiaand Newfoundland (see below).

The report of this species from GlacierNational Park, Montana by DeBolt &McCune (1993) is an error. A specimen ofthis collection (DeBolt 636) in WIS has asorediate thallus containing norstictic acidand a trebouxioid photobiont. Also in WISare several collections from Wisconsin ident-ified as F. praeruptorum, but these are F.recensa var. recensa (see below).

Fuscidea cyathoides var. suborientalis(Zahlbr.) M. Inoue, known only fromTaiwan and Japan also contains alectorialicacid (Inoue 1981a; as Fuscidea-1) and wasreported by Inoue as having curved, ellipsoidascospores of similar dimensions to thosereported for F. praeruptorum. The relation-ship between these two taxa is worthy of

further investigation, but, unfortunately, Ihave been unable to obtain any of Inoue’scollections for comparison because all theJapanese collections are presently unavail-able [on permanent loan from HIRO to Dr.Inoue (see below under F. scrupulosa)].

Selected corticolous specimens examined. Canada:British Columbia: Vancouver Island, along Pacific RimHwy, 22·5 km along the road S of Sutton Pass, 20 m onAlnus rubra, 1989, T. Tønsberg 12298b (BG). Newfound-land: Newfoundland Island, Avalon Peninsula NW,between Branch and North Harbour, 0–100 m, onAbies balsamea, 1991, T. Tønsberg 17103 (BG); BurinPeninsula, Placentia West District, c. 6 km ESE ofTerenceville, 54.6333(W, 47.6333(N, 150 m, 2000,C. Printzen 5668a (BG).—USA: Alaska: City andBorough of Juneau, Douglas Island E, West of Juneau,0·5 miles along Douglas Hwy NW of Juneau-DouglasBridge, 58(18#N, 134(27#W, 10 m, on Alnus rubra,1991, T. Tønsberg 16118 (BG); ibid., Baranof Island,c. 10 km E (direct) of Sitka, off the main road fromSitka to Herring Cove (Sawmill Cr. Rd) 60 m, on Alnusrubra, 1991, T. Tønsberg 16347 (BG; fertile).

Selected saxicolous specimens examined. Canada:Newfoundland: Newfoundland Island, Hawke Hills,47(19#20$N 53(07#37$W, 250–320 m, on rock,rocky, montane to subalpine oceanic heathland, 2007,J. Lendemer 10055, R. Harris 53845 (NY).—Sweden:Skaftö: Kristineberg, 14 August 1967, P. James s.n.(MSC).—USA: New York: Warrensburg, on rock inopen field, 1933, J. Lowe 3539 (MICH). West Virginia:Pendelton Co., Kile Knob, Fork Mountian,38(35#44$N, 79(28#57$W, 4588 ft [1398 m], onmassive sandstone, 2005, J. Vanderhorst 7119 (hb.Flenniken, MSC).

Fuscidea pusilla Tønsberg

Mycobank: MB358666.

Sommerfeltia 14: 138 (1992); type: Norway, Hedmark,Armot, Arset-Bechsminne, along state road 3, UTM32W PN 2674 (1917 II), 240 m., on Betula pubescens/pendula (road-side tree), 6 August 1983, Tønsberg 8041(BG—holotype; E, UPS—isotypes).

(Fig. 11)

Thallus usually of small thalli up to 10 mmdiam., composed of convex greenish areoles;prothallus distinct, pale to dark brown; soraliairregular, becoming confluent, soredia fari-nose; medulla I�.

Apothecia and conidiomata. Unknown.



Distribution and ecology. Fuscidea pusillawas first reported in North America byTønsberg (1993) from the AdirondackMountains, New York. It has subsequentlybeen collected in eastern North Americafrom Nova Scotia to West Virginia and inthe Pacific Northwest from WashingtonState to Kodiak Island, Alaska. This sug-gests that F. pusilla has an Appalachian-Great Lakes distribution in the east and anoceanic one in the west, but it appears tobe an infrequent species, at least in the east,and records are few. A collection from theThunder Bay District of Ontario on thenorthern shore of Lake Superior previouslyrefered to this species is better placed in F.recensa var. recensa.

Remarks. Fuscidea pusilla is the most fre-quent sorediate, corticolous species of thegenus containing divaricatic acid that hasbeen correctly reported from NorthAmerica. Fuscidea recensa var. recensa, whichis known from a single North Americancorticolous collection, differs in having amore widely-spread, predominantly brownthallus with discrete soralia, the outersoredia of which have a brown colouration.The European species Fuscidea lightfootii(Sm.) Coppins & P. James, which hasbeen incorrectly reported from the PacificNorthwest, is similar but has a morewidely-spread thallus, ulcerous soralia thatoften become confluent, and is usuallyfertile.

F. 11. Known North American distribution of Fuscidea pusilla. Includes some records from the Pacific Northwestand Newfoundland collected and identified by T. Tønsberg (BG) but not seen by the author.


Specimens examined. Canada: Nova Scotia: QueensCo., Kejemkujic N.P., Grafton Lake, along east shoreof Kejemkujic Lake near inlet from Grafton lake,44(23#N, 65(12#W, 110 m, hardwood forest(Quercus-Acer-Ostrya), on Betula papyrifera, 1999, I.Brodo 29675 (CANL); Cumberland Co., CapeChigneto Provincial Park, along trail from Red Rocks toMcGahey Brook, 45(21#00$N, 64(49–50#26–30$W,20–150 m, coastal forest, 2004, W. Buck 47134(NY).—USA: Connecticut: Litchfield Co., Town ofNorfolk, Holleran Swamp Preserve, E of Elmore Road,0·3 mi N of CT 272, 42(02#19$N, 73(12#23$W,hardwood-conifer swamp, 2003, R. Harris 47953 (NY).Maine: York Co., Town of Saco, Saco heath Preserve,43(32#48$N, 70(28#25$W, c. 60 m., Sphagnum-Ericaceae heath, with scattered trees of Pinus rigida andP. strobus, 2002, R. Harris 46314 (NY). New York:Duchess Co., Cary Arboretum, along E. BranchWappinger Creek, between Fern Glen and GiffordHouse, hardwoods, on Carya, 1981, R. Harris 14062(NY). West Virginia: Tucker Co., Monongahela N.F.,Olsen’s Bog, 39(07#N 79(36#W, 1000 m, open sprucebog, on twigs of shrub (Vaccinium) in bog, 2001, I.Brodo 30360 (CANL).

Fuscidea recensa (Stirt.) Hertel, V.Wirth & Vezda

Mycobank: MB342012.

Beitr. Naturk. Forsch. Südwestdeutschl. 31: 92 (1972).—Lecidea recensa Stirt., Scottish Naturalist 5: 219 (1880);type: Great Britain, Scotland, Perthshire, KinlochRannoch, Craeg Var, 09 1879, Stirton. (BM—holotype).

(Figs 12 & 13)

Thallus usually pale grey to brown andareolate, but occasionally almost white orred-brown and either continuous-rimose orareoles convex and bullate; soralia presentor absent, if present, discrete, initially palebut becoming brown, greenish whereabraded, soredia granular; medulla I�.

Apothecia black or brown, sessile, �con-stricted below, usually flat, 0·5–0·8 mmdiam. with a thin persistent proper exciple,often becoming flexuose and occasionallyalmost gyrose, occasionally convex andimmarginate; rarely pruinose. Thecium 55–65 �m tall, paraphyses simple, 1·5–2 �m,apical cell barely swollen to 3 �m, no pig-mented cap. Excipulum uniformly brownpigmented. Ascospores simple to 1-septate,ellipsoid, curved, 9–12�4–5 �m (Fig. 12).

Conidiomata not observed.


Distribution and ecology. Fuscidea recensa isa predominantly saxicolous species that iswidespread in eastern North America. Thenon-sorediate variety (var. arcuatula—seebelow), which is occasionally corticolous,is most frequent in the AppalachianMountains and the east coast, whereas thesorediate variety (var. recensa) is mostfrequent further west (Ozark Plateau,Wisconsin) and is almost exclusively saxi-colous, being known from a single NorthAmerican corticolous collection.

Remarks. Fuscidea recensa is characterizedby its long (10–12 �m), often curved, 0–1-septate ascospores (Fig. 12) and a thalluscontaining divaricatic acid. The type collec-tion and all European specimens are soredi-ate and only rarely fertile, whereas mostcollections referred to this species fromNorth America are fertile and lack soredia(e.g., Brodo et al. 2001). Fertile, non-sorediate collections are mostly from easternNorth America, whereas several collectionsfrom further west (Arkansas, Missouri and

F. 12. Fuscidea recensa var. arcuatula, ascospores of(Brodo 30610, Fryday 8873). Scale=10 �m.


Wisconsin) and one from West Virginia areboth sorediate and fertile (Fig. 13). Thefertile, non-sorediate specimens have anidentical apothecial anatomy and thallinechemistry to European collections and aretreated here as F. recensa var. arcuatula (seebelow).

Fuscidea recensa has an extremely vari-able thalline morphology, but all NorthAmerican collections with soredia andapothecia (i.e., var. recensa) have a thick,white thallus, whereas the thallus of non-sorediate collections is often dark grey.However, the thick, pale thallus morphologyis shared by non-sorediate specimens fromArkansas and North Carolina (see below)and also by several collections from thenorth-east (e.g., Brodo 22820B, Wong 435),whereas others (e.g., Brodo 30342 from WestVirginia) have an intermediate thick palebrown/fawn thallus. In fact, the thick, palethallus morphotype is the most frequent forthe species, specimens with a darker thallusoccurring in more exposed habitats, andthose with a thinner thallus in shaded ones.

Five other sorediate collections, fromIllinois, Kentucky, Ohio (two) and NewHampshire, lack apothecia but have aChlorella-type photobiont and are probablyreferable to F. recensa var. recensa, whereas acollection from North Carolina growingwith F. recensa var. arcuatula (Perlmutter 598p.p.), is definitely var. recensa. However,saxicolous sorediate crusts are poorly col-lected in North America, and it is probablethat F. recensa var. recensa is more wide-spread than reported here. The single NorthAmerican corticolous collection of var.recensa, from the Thunder Bay District ofOntario, has a shiny brown thallus withdiscrete soralia and and was previouslyidentified as F. pusilla.

Brodo et al. (2001) show the distributionof F. recensa as extending to northernColorado, but the specimen on which thisrecord was based is not a species of Fuscidea.In Europe, the sorediate taxon occasionallyoccurs on trees (Tønsberg 1992) but inNorth America var. recensa is almost exclu-sively saxicolous, although corticolous col-lections of var. arcuatula are relativelyfrequent.

Selected specimens examined. Great Britain: Scotland:V.C. 88 Mid Perthshire, Crieff, Ben Chonzie, 2250 ft,south facing crag, 1976, B. Coppins 2079 (E);Breadalbane Mts, Beinn nan Eachan, GR: 27/57.38,3000 ft, S-facing crags, exposed outcrop of schistoserock, 1985, B. Coppins (10997) et al. (E). V.C. 92

F. 13. Known North American distribution of Fusci-dea recensa. A, var. recensa (� soredia and apothecia, :

soredia only); B, var. arcuatula (●).


South Aberdeen, Ballochbuie Forest; above Bridge ofDee, GR: 37/18.90, 335–460 m, on vertical side of largeboulder on hillside, 1984, B. Coppins (10601) et al.(E).—USA: Arkansas: Stone Co., Cherokee WildlifeManagement Area, 35(45#46$N, 92(03#16$W, sand-stone bluff in mixed hardwoods, 2001, R. Harris 45464(NY), Buck 40351 (NY); Pope Co., Ozark NationalForest, Pedestal Rocks Scenic Area, 35(43#09$N,93(03#29$W, 550 m, sandstone bluffs in oak-dominated forest, 2005, W. Buck 49404 (NY); PolkCo., Ouachita National Forest, Caney Creek Wilder-ness Area, 34(24#18$N, 94(04#23$W, 1700 ft, oaks,hickory, and rock cliffs on S facing slope, 2000, C.Wetmore, 84444 (MIN). Missouri: Ste Genevieve Co.,Pickle Springs Natural Area, on boulder in shade, 1990,R. Harris 25986, 25992 (NY); Reynolds Co., St. Fran-cis Mountains, Johnson Shut-Ins State Park, 37(32#N,90(50#W, 245–310 m, oak-hickory-juniper forest onrhyolite and granite talus slope, 1993, R. Harris 31222(NY); Jefferson Co., St. Peters Sandstone ravine andglades, 38(23#13–41$N, 90(41#49$W, 165–260 m; onsandstone, 2001, W. Buck 45128 (NY). West Virginia:[Pocahontas Co.], Huntersville, 9 I 1930, F. Gray (F).Wisconsin: Portage Co., Cambrian sandstone, NWslope, 90(, shady and dry, 1962, KGF[oote] 62297(WIS); Juneau Co., Cambrian sandstone, level sunnyand dry, 1962, K. Foote 62696 (WIS); Vernon Co., St.Peter sandstone, NW slope, 90(, shady and dry, 1962,K. Foote 621096 (WIS); Clark Co., Cambrian sand-stone, S slope, 45(, sunny and dry, 1962, K. Foote61111a (WIS).

Sterile collections examined. Canada: Ontario:Thunder Bay District, Sleeping Giant P.P., west ofSilver Islet on trail to Sea Lion, 48(20#N 88(50#W,Abies-Picea forest, 1993, I. Brodo (29675) & S. Sharnoff(CANL; corticolous).—USA: Illinois: Union Co.,Panther Den, Shawnee National Forest, 37(33#N,89(05#W, moist sandstone bluff along river, maple-oakwoods and second growth, 1993, R. Harris 31432(NY). Kentucky: Harlan Co., Kentenia State Forest,Profile Rock, 2·7 mi SW of KY 2010 on Little ShepherdTrail, c. 765 m, large sandstone outcrop along ridge ofPine Mountain, 1991, W. Buck 20814 (NY). Ohio:Gallia Co., Wayne National Forest, above SymmesCreek, 8(49#34N, 82(28#19$W, 700–800 ft, south-west facing slope, on sandstone, 2006, J. Lendemer(7408, 7446) with participants of 15th TuckermanWorkshop (hb. Lendemer); Scioto Co., Niles Twp.Shawnee State Park, road to Lodge, 2006, D. Flenniken7703 with participants of 15th Tuckerman Workshop(hb. Flenniken). New Hampshire: Rockingham Co.,Nottingham, northern portion of North Mt,43(07#00$N 71(12#00$W, poor fen, volcano vent,erratics, mixed forest, 1994, on siliceous rock, E. Lay94-1588 (hb. Lay). North Carolina: Surry Co., PilotMountain State Park, 36(20#23$N 80(28#29$W, c.666 m, low elevation rocky summit plant community,sunny, 2006, G. Perlmutter (598 p.p.) & J. Pearson(MSC).

Fuscidea recensa var. arcuatula(Arnold) Fryday comb. nov.


Biatora arcuatula Arnold, Flora 71: 107–108 (1888).—Lecidea arcuatula (Arnold) Nyl. in Hue, Nouv. Archiv.du Muséum, sér 3: 129 (1891).—Biatora arcuatula (Arn.)Eckf., Bull. Torrey Bot. Club 22: 252 (1895).—Lecidearecensa f. arcuatula (Arnold) I.M. Lamb, Natl. Mus.Canad. Bull. 132: 254 (1954); type: Miquelon, 1884–1886, E. Delamare (W—isotype!, holotype not in Maccording to Hertel 1977).

Lecidea gyrodes H. Magn. in Degelius, Ark. Bot. 30A,no. 3: 36 (1942); type: USA, Tennessee, Great SmokyMountains, above Alum Cave, 1520 m, on granite rock,1939, G. Degelius (UPS—isotype!).

Remarks. Fuscidea recensa var. arcuatuladiffers from var. recensa only in lacking sor-alia and in being more or less confined toeastern North America.

Arnold (1887) initially determinedDelamare’s collection from Miquelon asBiatora circumflexa (Nyl.) Arnold, a speciesdescribed by Nylander from Lawrence Bayon the Asian side of the Bering Straits(Nylander 1885, 1887), but the followingyear described it as a distinct species, Biatoraarcuatula (Arnold 1888), specifically men-tioning the curved ascospores. Delamareet al. (1888) included the species underBiatora circumflexa, but Le Gallo (1952),although acknowledging Delamare’s et al.(1888) assertion that it is ‘‘commune surles rochers du Calvaire et du Capeau aMiquelon’’, mentions Lecidea circumflexa asdoubtfully present on Miquelon becauseZahlbruckner (1925) reported it only fromAsia.

Fuscidea recensa var. arcuatula, likeEuropean specimens of var. recensa, is veryvariable in thallus and apothecia gross mor-phology. The thallus varies from pale greyto dark brown, from thin and rimose to(usually) thick and areolate, whereas theapothecia are usually black and prominent,constricted below and with a thick persistentproper margin, but may also be paler, andalmost adnate with an �excluded margin,and occasionally with a slightly pruinosedisc. Lamb (1954) recognized the morphwith a dark brown thallus as f. arcuatula andBrodo (1984) suggested that this morph


may not occur in Europe because the typecollection of Lecidea recensa had a pale greythallus and ‘‘European authors . . . invari-ably describe the thallus as pale grey (Poelt& Vezda 1981, Wirth 1980)’’. How-ever, subsequent European authors havedescribed the thallus as pale grey to brown-grey (Purvis et al. 1992; Wirth 1995) orgreen to dark brown (Tønsberg 1992) and,in fact, European specimens of var. recensashow a similar variation in thallus colour tothose of var. arcuatula from North America;for example two collections from Scotland(Coppins 2079 & 10997) are as dark as theexsiccate specimen Brodo was describing(Gowan 5017) when he made this sugges-tion. Oberhollenzer & Wirth (1990) sug-gested that this variation in colour was aconsequence of the light intensity to whichthe specimen was exposed; specimens fromshaded habitats having a lighter thallus,whereas those from more exposed habitatswere darker. This is supported by specimensfrom North America; for example, twospecimens collected from the same area ofMaine (Brodo 30502, 30610), in which thespecimen from a shaded locality has a thin,pale thallus whereas that from an exposedrock has a thick, dark thallus. The speciesdescribed and illustrated by Brodo et al.(2001) as F. recensa, represents the darkform of this taxon.

The North American distributions of var.arcuatula and var. recensa are parapatric;Fuscidea resensa var. recensa having the centreof its North American distribution in centralUSA (Arkansas, Missouri, Wisconsin),where it is often fertile, with scatteredrecords of, usually, non-fertile, sorediatespecimens further east, whereas var. arcu-atula is largely confined to eastern NorthAmerica with only a few records from fur-ther west. However, the separation is notcomplete; in particular, three collectionsfrom western Arkansas (Polk Co.) are non-sorediate (i.e., var. arcuatula: Wetmore84283, 84371, 84572) whereas another col-lection from the same locality is morphologi-cally identical except for the presence ofsoralia (i.e., var. recensa: Wetmore 84444).The two taxa also occur together in the

southern Appalachians (Perlmutter 598) andthere is a record of F. recensa var. recensafrom New Hampshire (Lay 94-1588). It isprobable that the overlap may be more ex-tensive because sorediate saxicolous crustoselichens are infrequently collected in NorthAmerica, and non-fertile var. recensa may bemuch more common in the east than recordsindicate. For this reason I prefer, at thisstage, to recognise the non-sorediate entityat the level of variety rather than as a distinctspecies.

The type, and only collection of Lecideagyrodes H. Magn. differs from typical F.recensa var. arcuatula only in having a thalluscomposed of brown, convex areoles andsomewhat gyrose apothecia, other apothecialcharacters being identical to typical F.recensa. These gross morphological charac-ters are at the extreme range of the morpho-logy of the taxon, but specimens with anequally dark thallus or consisting of equallyconvex areoles are known. For example, acorticolous collection from West Virginia(Lay 01-0460) has a thick, �bullate thallusthat is partly dark brown (as in L. gyrodes)and partly pale, but has non-gyrose apoth-ecia. Contorted, �gyrose apothecia are,however, known from some specimens oftypical var. arcuatula and, consequently, L.gyrodes is here included as a synonym of F.recensa var. arcuatula.

Fuscidea ramboldioides Kantvilas (Kantvilas2001, 2004), known from Australia (NewSouth Wales, South Australia, Tasmania),has similar curved 0–1 septate ascosporesbut these are shorter than in F. recensa and italso differs in having a smoother thallus anda less pigmented exciple.

Selected specimens examined. Canada: New Brunswick:Albert Co., Fundy N.P., Matthew’s Head, 45.5667(N64.9667(W, in old field on unshaded boulder, 1981,S. Gowan (5017) & I. Brodo (DUKE, MSC, WIS,Lichenes Canadenses Exsiccati #178). Newfoundland:Labrador, L’anse au Clair, 1894, A. Waghorne 20, p.p.(US); Sparrible Cove, Notre Dame Bay, 1894, A.Waghorne 23, (MIN); Newfoundland Island, AvalonPeninsula, Hawke Hills, 47(19#20$N 53(07#37$W,250–320 m, on rock, rocky, montane to subalpineoceanic heathland, 2007, J. Lendemer 10011, 10050(NY), E. Lay 07-0025 (hb. Lay); ibid., ButterpotProvincial Park, E of Big Otter Pond, 47.399(N53.043(W, 160 m, mesic to wet coniferous Pinus


mariana forest, rock bluffs along banks, 2007, E. Lay07-0026, 07-0028, 07-0029 (hb. Lay). Nova Scotia:Inverness Co., Cape Breton Highland NP, Presqu’ile,46(42#N 60(57#W, on exposed rocks on bluff, 1972, I.Brodo 18979 (CANL, WIS); ibid., Cape Breton Island,Cape Breton Co., Mainadieu, on cliff face about 10 mabove the sea, 1952, I. Lamb 6916 (CANL). Ontario:Leeds & Grenville Co., Morton, Pitch Pine Ridge,44(24#N 75(53#W, on granite cliff, 1968, P. Wong435 (CANL).—USA: Arkansas: Polk Co., OuachitaNational Forest, Caney Creek Wilderness Area,34(25#22$N, 94(08#22$W, 1150 ft, N facinghillside and river bottom, 2000, C. Wetmore 84283(MIN). Maine: Knox Co. Rockport, on quartz rock, 9 v1911, G. K. Merrill, (FH, Lichenes Exsiccati #256);.Washington Co., McLellan Park, 44(29#N 67(51#W,on exposed rocks at edge of shore, 2001, I. Brodo(30583), F. Brodo & F. Schumm (CANL); Steuben,Eagle Hill Field Research Institute, 44(28#N67(56#W, under overhang, in crevice of rockface, 2001,I. Brodo 30502 (CANL); ibid., open disturbed site,44(29#N 67(56#W, on exposed boulder, 2001, I.Brodo 30610 (CANL); Town of Beals, Great WassIsland, Great Wass Island Preserve, 44(28#52$N67(35#41$W, red spruce-balsam fir coastal forest andjack pine bald, 2007, R. Harris 53803 (NY); HancockCo., Black Mountain trail, just south of Tunk Lake.44(34#50$N 68(6#22$W, c. 230 m, Picea rubens stand,on boulder on trail in forest, 2004, I. Brodo (31488) &Eagle Hill class (DUKE); ibid., Black Mountain,44(34#52$N 68(6#24$W, c. 246 m, on boulder in oldmaple forest, shady, 20 vii 2006, G. Perlmutter s.n.(MSC). Massachusetts: Essex Co., Cape Elizabeth,1861, E. Tuckerman (FH); New Bedford, 1862–1898,H. Willey (FH, US), Weymouth, 1862–1898, H. Willey(US); Dukes Co., Martha’s Vineyard, Francis WoodTNC Preserve, 41(24#N 70(42#W, old field, on rock-fall, 1994, S. D. Sharnoff & S. Sharnoff 1511.01(CANL–LNA voucher). New Hampshire: Coos Co.,Great Gulf Wilderness Area, around Great Gulf Trailand W. Peabody River, 2300 ft, along ridge and river,1988, C. Wetmore 61838, 61842 (MIN); WhiteMountains, 1843, E. Tuckerman (FH); Mt Monadnock,C. Frost (FH); White Mountains, 1878, H. Willey (US).New York: Washington Co., West Fort Ann, flinty rockseast of Lake Hadlock, 24 xi 1915, S. Burnham (FH, exhb. G.K. Merrill); Essex Co., Wilmington, notch on Rt.86, 5·8 miles S of Wilmington, exposed talus slope onsteep grade, 1963, I. Brodo 4714 (CANL, MSC). NorthCarolina: Surry Co., Pilot Mountain State Park (MtnSection), 36(20#23$N 80(28#29$W, c. 666 m, lowelevation rocky summit plant community, sunny, 2006,G. Perlmutter (594, 598) & J. Pearson (MSC); 1883,Jos. Hooker (FH, US); ibid., Crowden Mt, 1883, H.Green (PH); ibid., 1886, H. Green (US). Pennsylvania:Bedford Co., Buchanan State Forest, CumberlandValley township, Pleasant Valley, 40(03#18$N,78(39#43$W, 1700–1800 ft, west facing slope withgranite exposures, on sandstone, 2006, J. Lendemer7285 (with W. Buck, R. Harris, S. LaGreca & N. Wood)(hb. Lendemer). South Carolina: Mitchell Co., RoanMountain, Roan High Bluff, 6200 ft, spruce-fir, on

Picea, 1895, R. Harris 18305 (NY: no apothecia, divari-catic acid); Aiken, H. Ravenel (FH); Chester, 1883, J.Eckfeldt (PH). Tennessee: GD[egelius] (MIN; ex hb. G.Llano; topotype of L. gyrodes; possible isotype); ibid.,Great Smoky Mountains, near Alum Cave, c. 1500 m.13 ix 1939, G. Degelius (US—topotype of L. gyrodes;possible isotype). Vermont: Addison Co., Bristol, BristolCliffs Wilderness Area, 44(06#19$N 73(04#34$W,cold air talus slope, silicous, 2000, E. Lay 00-0071 (hb.Lay); Lamoille Co., Battleboro & Mt Mansfield, C.Frost (FH); Mt Mansfield State Forest, Runny NoseSpring, 44(31#41$N 72(48#54$W, krumholtz and sili-ceous ledges in open, 2000, E. Lay 00-0072 (hb. Lay).Virginia: Bedford Co., The Peaks of Otter, 3875 ft,oak-hickory forest over granite and sandstone, on rockoverhang, 1978, I. Brodo (22820B) & D. Pierson(CANL). West Virginia: Grant Co., Greenland Gap,39(11#07$N, 79(08#25$W, 450–475 m, on rocksat edge of dashing brook, 2001, S. LaGreca 799(FH); Tucker Co., Monongahela N.F., Dolly SodsWilderness, Bear Rocks, 39(04#N 79(18#W, 1217 m,on boulder on ridge, 2001, I. Brodo 30342 (CANL);Pendelton Co., Kile Knob, Fork Mountian,38(35#44$N, 79(28#57$W, 4588 ft [1398 m], onmassive sandstone, 2005, J. Vanderhorst 7119 (hb.Flenniken).

Corticolous or lignicolous. USA: Maine: WashingtonCo., Eagle Hill, 44(27#30$N, 67(56#00$W, Acerrubrum, 2006, R. Harris 53102 (NY). New York: UlsterCo., Ellenville Ice Caves, 1937, collector unknown(NY, ex Herb. G. G. Nearing). Vermont: Addison Co.,Bristol, Bristol Cliffs Wilderness Area, 39(04#34$N44(06#19$W, cold air talus slope, siliceous, 2000, E.Lay 00-0071 (hb. Lay). West Virginia: Tucker Co.,Monongahele National Forest, Dolly Sods Wilderness,near headwaters of Red Creek, on Acer rubrum, 1975, R.Harris 10446 (NY); ibid., Bear Rocks, exposed stuntedtrees, 2001, E. Lay 01-0460 (hb. Lay).

Fuscidea scrupulosa (Eckf.) Frydaycomb. nov.


Biatora scrupulosa Eckf., Bull. Torrey Bot. Club 22: 253(1895).—Lecidea (Biatora) scrupulosa (Eckf.) H. Magn.,Meddel. Göteborgs Bot. Trädgard/Acta Horti. Gothob. 10:9 (1935); type: Canada, Labrador, on rock, 1894, A. C.Waghorne (PH—lectotype!, designated here).

Lecidea kochiana var. subreagens H. Magn. syn nov.,Meddel. Göteborgs Bot. Trädgard/Acta Horti. Gothob. 10:10 (1935).—Fuscidea subreagens (H. Magn.) Oberholl.& V. Wirth, Beiheft zur Nova Hedwigia 79: 575 (1984);type: USA, New York, Adirondak Region, MountMarcy, 4900–5300 ft, 1933, J .L. Lowe 2701 (UPS—holotype), 2836, 3111, 3096 (FH, NY, US—syntypes!).

(Figs 14 & 15)


Thallus grey to brown (becoming reddish-orange in the herbarium), moderately thick,areolate, areoles flat, thin black prothallus atmargin; medulla I�.

Apothecia black to dark-brown, rounded,becoming slightly flexuose, 0·5–0·9 mmdiam.; proper margin persistent and slightlyraised, 0·05–0·1 mm wide. Thecium 70–80 �m tall; paraphyses sparingly branchedwith upper 5 �m pigmented, apical cellsswollen to 5 �m with a brown cap. Exciple ofradiating hyphae, outer 20 �m pigmented,inner colourless. Ascospores simple, becom-ing brown, sub-globose to broad-ellipsoid,8–10�5–6 �m

Conidiomata pycnidia, rare, immersed inthallus; conidia bacilliform 3–4�0·8 �m.

Chemistry. K�, C+ red, KC+ red, Pd+yellow, UV+ orange. Alectorialic acid byTLC.

Distribution and ecology. This saxicolousspecies is confined to higher altitudes innorth-east North America (Fig. 14) where itis not rare.

Remarks. Fuscidea scrupulosa is character-ized by the presence of alectorialic acid inthe thallus and apothecia. This substance isresponsible for the ‘yellow mist’ producedwith K that was noted by Magnusson(1935), and also causes the thallus to reactPd+ yellow, KC+ red, UV+ orange.Alectorialic acid is otherwise known in thegenus in North America only in the sorediatespecies F. praeruptorum (Du Rietz & H.Magn.) V. Wirth & Vezda.

It is surprising that Magnusson (1935) didnot realize that his new variety, Lecideakochiana var. subreagens, was identical to thespecies he newly combined into the Lecidearivulosa group in the same publication. Hedescribed them in successive entries andkeyed them out together in the samecouplet, separating them by L. scrupulosahaving a thicker thallus, sessile apotheciawith a more prominent margin, and slightlysmaller ascospores. However, it is probablethat Magnusson had seen only the holotypematerial of his new variety sent to him byLowe and the fuller range of Lowe’s collec-tions from Mt Marcy, and numerous othercollections from NE North America, showthat these distinctions are not consistent.

Fuscidea circumflexa (Nyl.) V. Wirth &Vezda, described from the Asian side of theBering Straits, is very similar to F. scrupulosaand may be conspecific with it, as it appar-ently differs only in having a thicker,verrucose thallus (Fig. 15). The lectotype ofF. circumflexa (not holotype as stated byInoue 1981a) is extremely poor, being lessthan 1 cm2 and having only two apothecia(both of which have been sectioned by pre-vious investigators). An isotype in S is larger,although still small, and has several apoth-ecia, but it is impossible to be certain thatthe two species are not conspecific. Conse-quently, given that the two taxa are allopat-ric and appear to differ in gross morphology,I here retain them as separate species. Asimilar situation occurs with Lecidea gyrodesand F. recensa var. arcuatula (see above), butin that case other specimens of F. recensa var.arcuatula exist with the properties of L.gyrodes and, as the two entities are not widelyseparated geographically, they are included

F. 14. Known distribution of Fuscidea scrupulosa.


in the same taxon. Inoue (1981a) reportedF. circumflexa from Japan but from hisdescription ‘‘areoles plane or slightly con-vex’’ and his illustration, it appears hisspecimens may represent F. scrupulosa.Examination of these collections may help todetermine whether F. scrupulosa should bereduced to synonymy with F. circumflexa,

but, unfortunately, I have been unable toobtain any for study because they are onpermanent loan from HIRO to Dr Inoue.

Eckfeldt’s collection of F. scrupulosa in PHis marked ‘syntype’, but I have seen no otheroriginal collections among the numerousother collections I have examined fromall the major North American herbaria.

F. 15. Thallus and apothecia of A, Fuscidea scrupulosa (holotype) and B, F. circumflexa (isolectotype). Scales: A& B=5 mm.


Eckfeldt mentions no specific collectionsand the PH specimen is an excellent speci-men in good condition so I here designate itas the lectotype.

Selected specimens examined. Canada: Newfoundland:Labrador, L’anse au Clair, 23 vii 1894, A.C. Waghorne(US, p.p., not type material); Newfoundland Island,Avalon Peninsula, Hawke Hills, 47(19#20$N53(07#37$W, 250–320 m, on rock, rocky, montane tosubalpine oceanic heathland, 2007, J. Lendemer 10011,10057 (NY). Quebec: N side of Lake Louis, c. 80 mi Nof Schefferville, on rock, 1990, D. Wehr 39 (NY).—USA: Maine: Hancock Co., Acadia N.P., Mt DesertIsland, in gorge, birch, oak and young spruce woods,1984, T. Sullivan 4386 (MIN); Piscataquis Co., BaxterState Park, Katahdin, North Basin, 45(55.76#N,68(54.86#W, 975 m, granitic boulder, 2004, Fryday8843, 8844 (MSC); ibid., Tableland, 18 ix 1922, A.Norton, Pope & Perkins (FH). New Hampshire: CoosCo., Great Gulf Wilderness Area, NE slope of Mt Clay,6050 ft, in alpine area with rock tallus, heath meadowand some areas of krumholtz, 1988, C. Wetmore, 61623(MIN); ibid., Presidential Dry River Wilderness Area,on east side of Mt Monroe, 5000 ft, in sedge and heathmeadow in alpine area, 1988, C. Wetmore 62426(MIN); White Mountain NF, Mt Washington, alongAuto Road at Cragway Spring, 44(17#N, 71(18#W,1430–1465 m, heath and krummholz vegetation onleeward slope, 1999, R. Harris 4319 p.p. (NY), W. Buck36167 (NY); White Mountains, 1843, E. Tuckerman(FH); Mt Monadnock, C. Frost (FH). New York: EssexCo., Mount Marcy (near Lake Placid), 5300 ft, on rockon N side, 1933, J. Lowe 2604 p.p. (MICH); ibid.,underside of rock NE exposure, 1933, J. Lowe 2607p.p. (MICH); ibid., Avalanche Lake (near Lake Placid),1000 ft, 1936, J. Lowe 6237 (MICH); Lake Placid,Algonquin Peak south of Mt Marcy, 5110 ft [1558 m],1963, I. Brodo 4750 p.p. (CANL, MSC); Wilmington,Whiteface Mountain, summit, c. 4500 ft, on easternslope in krumholtz zone, in shady little nook, 1960, I.Brodo 492 (CANL, MSC); ibid., 4872 ft, on top abovetree line, 1963, C. Wetmore 12209 p.p. (MIN); ibid., atsummit, on rocks, 1963, J. Thomson 11221, 11235(WIS); ibid., near summit of Whiteface Mountain, 8miles NE of Lake Placid, 44(22#N, 73(54#W, 4800 ft(1445 m), Abies balsamea krumholtz, 1971, R. EganEL-4678, EL-4680 (MIN). Vermont: C. Frost (FH);Lamoille Co., Mt Mansfield State Forest, Runny NoseSpring, 44(31#41$ 72(48#54$W, krumholtz and sili-ceous ledges in open, 2000, E. Lay 00-0073, 00-0074(hb. Lay).

Fuscidea texana Fryday sp. nov.


Fuscideae molli similis sed thallo acidum norsticticumcontinenti differens.

Type: USA, Texas, Brewster Co., Big Bend NationalPark, on north slope of Emory Peak, [c. 29(15#N,

103(15#W,] 6300 ft, pinyon pine, oaks and junipers, 16May 1970, C. M. Wetmore 19651 (MIN—holotype!).

(Figs 10 & 16)

Thallus to 5 cm diam., with distinctmargin, c. 0·2 mm thick, pale pinkish-grey,areolate; areoles flat, 2·5–4·0 mm across,often subdivided into smaller ‘subareoles’;cortex absent but with numerous minutebrown crystals on the surface; medulla I�.Photobiont Chlorella–type, cells 8–12(–15when dividing) �m diam.; dividing by binaryfission.

Apothecia sessile to adnate, (0·3)0·6–1·0(1·3) mm diam. mostly round but someirregular, dark brown, slightly pruinose;margin indistinct even in very young apoth-ecia, when present barely discernable as athin, pale margin 0·01–0·02 mm across andnot raised above the level of the disc.Thecium 50–60 �m tall; paraphyses simple,lax and readily separating in K, 2–2·5 �mwide with upper 10–15 �m with brown pig-ment, distinctly capitate, to 5 �m across withdark brown pigmented cap. Hypotheciumhyaline 75–80 �m tall. Ascus Teloschistes-type, 40–55�10–12 �m, cylindrical toslightly clavate; ascospores broad-ellipsoid,8–9�4·5–5·5 �m, thick walled, becomingpale brown with age. Exciple cupular, verythin with only the outer 5 �m with brownpigment.

Conidiomata rare, immersed, pale reddish-brown 0·05 mm diam.; conidia narrow ellip-soid 5–6�1·5–2·0 �m.

Chemistry. K+ red (acicular crystals insection), C�, Pd+ yellow; norstictic acid.

Distribution and ecology. The occurrence ofa species of Fuscidea in southern Texas isunexpected. The genus otherwise exhibits apredominantly cool temperate-boreal/australdistribution, being especially frequent inareas with an oceanic climate. SouthernTexas is in the ‘‘arid-subtropical’’ climaticregion, and although F. texana was collectedfrom high elevations (1920 and 2255 m), theclimate is still dry with an annual precipita-tion of 48·7 cm, and quite unlike that of any


other area where the species of the genusoccur in North America. However, Fuscideafulva (Malme) Kalb from Central and SouthAmerica, F. hottentotta Brusse from SouthAfrica, F. umbricolor (Nyl.) Hertel fromnorthern South America, and an unde-scribed species from the West Indies (A. M.Fryday unpublished.) also do not fit withthis general distribution pattern. Thesespecies differ from F. texana in chemistryand, in addition, F. fulva is corticolous, F.hottentotta has curved ascospores, and F.umbricolor has an amyloid medulla.

Remarks. Norstictic acid is otherwiseknown in Fuscidea only in two southernhemisphere species; F. subasbolodesKantvilas from the cool-temperate/subantarctic (Kantvilas 2001; Fryday 2003)and F. mayrhoferi Kantvilas from WesternAustralia (Kantvilas 2004). These speciesdiffer in having innate apothecia with an‘intercincta-type’ exciple (Oberhollenzer &

Wirth 1984). The conidia of F. texana arelonger than is usual in the genus, mostother species having conidia 2·5–4�1·2–2·0 �m, although those of F. mayrhoferi arereported as 4–5�1–2 �m (Kantvilas 2004)and in the related Ropalospora lugubris(Sommerf.) Poelt they are bacilliform,4–6�1 �m.

Only nearly 40-year-old herbariummaterial of F. texana was available for studyand given that the thallus apparently lacks acortex, the phobiont cells reaching to theupper surface of the thallus, this species islikely to appear a very different colour whenfresh.

Other lichen species associated with theholotype collection of F. texana includedCaloplaca sp., Candelariella sp. and Dime-leana sp.

Additional specimen examined. USA: Texas: BrewsterCo., Big Bend National Park, on top of east rim,[29(14#N, 103(17#W], 7400 ft, in pinyon pine-juniper

F. 16. Thallus and apothecia of Fuscidea texana (Wetmore 19651, holotype). Scale=1 mm.


forest and on edge of cliffs, 1969, C. Wetmore 18333(MIN, MSC).

Fuscidea thomsonii Brodo & V. Wirth

Mycobank: MB446213.

Lichenographia Thomsoniana: 156 (1998); type: Canada,British Columbia, Queen Charlotte Islands, MoresbyIsland, east end of Mike Inlet on shore and along streamto exposed bluffs, 52(32#N 131(45#W, 36 m, 1 Aug.1968, I. M. Brodo 14173 (CANL—holotype).

(Figs 1E, 2 & 3)

Thallus grey to greyish-brown, crackedareolate; black prothallus usually present atmargin; medulla I+ violet.

Apothecia immersed to adnate, not con-stricted below; disc black, flat; pseudo-thalline margin usually apparent, paler thanthe disc giving the appearance of a ‘halo’intercincta-type (cf. F. aleutica). Thecium 55–75 �m; paraphyses simple, lax in K, upper20–25 �m with brown pigment, 1·5–2·0 �mwide scarcely swollen at apices (to 3·0 �m).Exciple annular, poorly defined, only outer-most cells with brown pigment. Ascosporessubglobose to broad-ellipsoid, 7–9�5–6 �m.

Conidiomata pycnidia, immersed in thal-lus; conidia bacilliform 3–4�0·8 �m.


Distribution and ecology. This species wasdescribed from the Queen Charlotte Islandsby Brodo & Wirth (1998) and is here re-ported for the first time from Alaska fromtwo collections from the Aleutian Islandspreviously identified as F. lowensis (Talbotet al. 1997).

Remarks. Fuscidea thomsonii is character-ized by having innate apothecia and a palegrey thallus containing divaricatic acid andan amyloid (I+ violet) medulla. Fuscideaaleutica is similar but differs in having abrown thallus, more sessile apothecia with abetter developed exciple that lack a whitering around the inner margin, and a more

densely pigmented epihymenium andexciple.

Two collections with a dispersed thallus,which were mentioned by Brodo & Wirth(1998), are provisionally included here (seebelow). In one instance, the two forms occuron the same collection (Brodo 26977) andadditional material may show that they rep-resent distinct taxa. One collection (Brodo26977) considered by Brodo & Wirth (1998)to be an aberrant form of F. mollis, is hereincluded in F. thomsonii, whereas threeother collections from the Queen CharlotteIslands (Brodo 10444 p.p., 10455, 17833),mentioned by Brodo & Wirth (1998) aspossibly an unnamed species or aberrantforms of F. thomsonii, are included in F.aleutica.

Selected specimens examined. Canada: BritishColumbia: Queen Charlotte Islands, Graham Island,Port Channal off Athlow Bay, on south side of inlet(Goose Cove), shoreline rocks and trees and bluffs, onboulder at summit, 320 ft, 1967, I. Brodo (10444 p.p.),M. Shchepanek & W. Schofield (CANL); ibid., MoresbyIsland, on west coast, Kootenay Inlet, spruce forest andPinus contorta-Chamaecyparis stand on bluffs, 52(51#N132(13#W, on top of exposed bluff, 1967, I. Brodo(12155) & M. Shchepanek (CANL); ibid., HibbenIsland (off west coast of Moresby Island), 53(00#N132(22#W, headland rocks and rocky shore at northend of island, on rock in salt-spray zone, 1968, I. Brodo14058 (CANL); ibid., Chaatl Island, cove on westcoast, on NE corner of island, facing ocean, 53(08#N132(35#W, 0–9 m, on lower aerohaline on headlandrocks, 1988, I. Brodo 26977 p.p. (CANL); ibid.,Ingraham Point, entrance to Carpenter Bay, southshore, 52(14#N 131(02#W, exposed headland rocks,,on rocks in salt-spray zone, 2000, I. Brodo 29958(DUKE, MSC, WIS: Lichenes Canadenses Exsiccati#260), 29959 (CANL).—USA: Alaska: Aleutianislands, Adak Island, Finger Bay area, 51(49.782#N,176(37.886#W, 0·5 m, on coastal outcrop, epilithic,1994, S. & S. Talbot 635 (WIS); ibid., 51(49.639#N,176(37.634#W, 200 m, alpine tundra, on rocks in creekbed, 1994, S. & S . Talbot 652A (WIS).

Specimen with dispersed thallus examined. Canada:British Columbia: Queen Charlotte Islands, ChaatlIsland, cove on west coast, on NE corner of island,facing ocean, 53(08#N 132(35#W, 0–9 m, on loweraerohaline on headland rocks, 1988, I. Brodo 26977p.p. (CANL): ibid., Graham Island, Seal Inlet inRennell’s Sound, bluffs and boggy area above forest andshoreline around a small cove on the west shore of theinlet, 53(31#N 132(44#W, on vertical faces of exposedrocks, 1967, I. Brodo (10455) & M. Shchepanek(CANL).


Excluded species

Fuscidea cyathoides (Ach.) V. Wirth &Vezda

Mycobank: MB417673.

Beitr. Naturk. Forsch. Südwestdeutschl. 31: 92 (1972).—Lichen cyathoides Ach., Lichenogr. Suec. Prodr.: 62(1798); type: Sweden, in saxis et rupibus (H-Ach 273 F& G—lectotype, fide Oberhollenzer & Wirth 1984:552).

Fuscidea cyathoides is a saxicolous specieswith bean-shaped ascospores and a thalluscontaining fumarprotocetraric acid (Pd+red). The occurrence of this species in NorthAmerica was discussed by Thomson (1997)who concluded that it has not been reliablyreported from the continent; a conclusionwith which I fully concur.

Specimens examined. France: Vosges: Col de laSchlucht, 1140 m, sur Fagus, 25 9 1959, R. Werner(MSC).—Great Britain: Scotland: V.C. 92 SouthAberdeenshire: Braemar, Invercauld Estate, CraigLeek, 37(NO)/192.931, 400 m, quartzite outcropsbelow crags, 2005, Fryday 9011, 9012 (MSC).—Norway: Nordlandia: Grolo, J. Norman (DUKE).

Fuscidea kochiana (Hepp) V. Wirth &Vezda

Mycobank: MB342006.

Beitr. Naturk. Forsch. Südwestdeutschl. 31: 92 (1972).—Lecidea kochiana Hepp, Lichenen Flora von Würzburg: 61(1824); type: Germany, Hessen, Rhon, Milsburg,770 m, 21 ix 1968, V. Wirth (STU-WIRTH 885—neotype, fide Oberhollenzer & Wirth 1984: 570).

(Fig. 1F)

Fuscidea kochiana is a saxicolous specieswith innate apothecia and a thallus contain-ing divaricatic acid. It was reported fromNewfoundland by Eckfeldt (1895) and fromthe Adirondacks (New York) by Lowe(1939). However, no specimens of thisspecies could be found in PH, where Eck-feldt’s collections are housed (J. Lendemerpers. comm.) and Eckfeldt’s description ofthe thallus as ‘‘pale ashy brown to fuscesent,rimose, areolate, broken, areoles sunken,plane, bordered by a black margin’’ suggeststhat his specimen is not this species. Lowe’scollections (in MICH) determined as Fusci-

dea kochiana by Lowe and Magnusson arereferable to other species of Fuscidea (mostlyF. appalachensis and F. scrupulosa). Fuscideakochiana was typified by Oberhollenzer &Wirth (1984) and Magnusson appears tohave had a different concept of this speciesfrom that now accepted. Although he cor-rectly identified the distinguishing morpho-logical character as the innate, immarginateapothecia (Magnusson 1925), he deter-mined several of Lowe’s collections from theAdirondacks as this species, even thoughthey had sessile apothecia with a well-developed margin. He also described thenew variety F. kochiana var. subreagens (asynonym of F. scrupulosa, see above), whichhas sessile apothecia with a well-definedmargin, as ‘‘outwardly not differing from thetype which is collected in the same locali-ties’’ (Magnusson 1935). Fuscidea kochianawas not treated by Thomson (1997) and as Ihave seen no other collections from NorthAmerica I conclude that Fuscidea kochianahas not been correctly reported from thecontinent.

Fuscidea kochiana was also recentlyreported from Brazil (Aptroot 2002) butthe collection upon which that report wasbased is referable to a species similar toF. umbricolor, but lacking an I+ medulla(cf. Additional comparative materialexamined—Materials and Methods).

Specimens examined. France: [Vosges:] La Schlucht,1890 (Harmand, Lichenes in Lotharingia, #1046)(MSC).—Great Britain: Scotland: V.C. 92 SouthAberdeen: between Meall Odhar and Glas Maol, 37/1577, 900 m, on top of siliceous rock outcrop (?schist),1995, Fryday 6073 (herb. Fryday); ibid., Braemar,Eastern Cairngorms SSSI, Mar Lodge Estate, ClaisFhearnaig, 37(NO)/066.930, 505 m, acid rock boulderfield above lochan on S side of narrow valley, 2005,Fryday 8978 (MSC).

Fuscidea lightfootii (Sm.) Coppins &P. James

Mycobank: MB342007.

Lichenologist 10: 201 (1978).—Lichen lightfootii Sm, inSowerby, English Botany 21: tab. 14541. 1805; type:Great Britain, England, Sussex, St. Leonards Forest,W. Borrer (BM—holotype).


Fuscidea lightfootii is a corticolous, soredi-ate species with a thallus containing divari-catic acid, and ‘dumbell-shaped’ ascospores(cf. Kantvilas 2002; fig. 1C). It was reportedfrom British Columbia by Aptroot (1996),but the two collections upon which thisreport was based were shown to be refer-able to other species by Tønsberg (2002).Collections from the north-east previouslyreferred to this species are F. arboricola.

Fuscidea subfilamentosa (Zahlbr.)Brako

Mycobank: MB132216.

In Egan, Bryologist 90: 163 (1987).—Phyllopsora subfila-mentosa Zahlbr., Ann. Mycol. 33: 44 (1935); type: USA,Florida, [Seminole Co.], Sanford, on dead trunk ofQuercus virginiana, 3 10 1919, S. Rapp (W—holotype!).

This species was described from a collec-tion from Florida that is, to my knowledge,still the only record. The holotype in W, aspecimen in F, and collections from Rapp’sherbarium in FLAS (not seen) all appear tobe from the same collection, having beencollected by S. Rapp from the same localityin March 1919, the only difference beingthat the holotype has the number ‘‘62’’ andthe name ‘‘Psora’’ whereas the F collectionand one of the FLAS collections have thenumber ‘‘3109’’ and the name ‘‘Phyllopsorasubfilamentosa’’. The other FLAS collectionhas no number and the name ‘‘Phyllopsorasubfilamentosa’’ (R. Harris, pers. comm.). Itis unlikely that these are collection numbers,as this would have meant Rapp would havecollected almost 3000 collections in a singlemonth, but are more likely numbers refer-ring to individual determinations or taxa. Itappears that Rapp initially determined thecollection as Psora and gave it the number62; he then sent part of the collection toZahlbruckner, who, 16 years later in 1935,named it Phyllopsora subfilamentosa, at whichpoint Rapp renamed his duplicate and gaveit the new number 3109. Unfortunately,although the FLAS and F collections areclearly topotypes, it is not possible to posi-tively identify them as isotypes.

The holotype in W is in good conditionwith numerous apothecia, but it is not a

species of Fuscidea. It is easy to see, however,why this species was referred to that genus.The apothecia have exclusively brownapothecial pigments and simple hyaline as-cospores that soon become pigmented. Onthe other hand, the pigment is more red-brown than the fuscous-brown in Fuscidea,the ascospores are larger (14–16�8–9 �m)than those of any known Fuscidea sp., andlarge, dark blue (K+ bright blue, 15%HCl+ violet) granules are present in thehypothecium. In addition, the specimen hasa squamulose thallus, and the exciple iscomposed of swollen radiating hyphae—both characters unknown in Fuscidea. More-over, the ascus apex has an IKI+ blueconical ‘plug’ or ‘tube’ of the Porpidia-typeand is clearly not Teloschistes-type.

All the above characters suggest that thisspecies is referable to the Lecidea hypnorumgroup. This group is closely related toClauzadea and is not congeneric with thetype species of Mycobilimbia, where it isoften placed. It is worthy of generic rank butas no generic name is currently available thespecies is here transferred to Lecidea and thenecessary new combination made.

Lecidea subfilamentosa (Zahlbr.)Fryday comb. nov.


Phyllopsora subfilamentosa Zahlbr., Ann. Mycol. 33: 44(1935); type: USA, Florida: [Seminole Co.], Sanford,on dead trunk of Quercus virginiana, 3 10 1919, S. Rapp(W— holotype!).

Thallus composed of minute, dispersed,slightly convex, pale, creamy grey squam-ules, 0·4–0·7�0·15 mm. Photobiont chloro-coccoid, cells 5–6(–8) �m diam.

Apothecia lecideine, 0·4–0·6 mm diam.,flat, constricted below, with a black to red-brown disc and black, slightly raised propermargin 0·05–0·1 mm wide. Hymenium 70–80 �m tall, epihymenium 15 �m tall, diffusepale golden-brown (superba-brown); para-physes, slender, 0·1–0·15 �m thick, simple,conglutinate, slightly branched and septateabove, constricted at septa, apical cells�swollen to 3 �m. Hypothecium goldenbrown, K+ brown (superba-brown). Exciple


T 1. Key characters of Fuscidea species reported from North America (italics) and outside North America (no italics)

Taxon Substratum Distribution Soredia Chemistry

Thallus

Apothecia AscosporesColour and texture Medulla

F. aleutica saxicolous western absent divaricatic acidUV+ white

brown, thin, rimose I+ violet sessile broad-ellipsoid

F. appalachensis saxicolous eastern �present divaricatic acidUV+ white

grey, areoles convex I� sessile subglobose

F. arboricola corticolous eastern present fumarprotocetraric acidPd+ red

grey I� sessile straight

F. circumflexa saxicolous Japan(?),BeringStraits

absent alectorialic acidC & KC+ red;Pd+ yellow UV+ orange

brown, areoles convex I� sessile straight

F. cyathoides saxicolous Europe,Asia

absent fumarprotocetraric acidPd+ red

brown or grey, smooth I� sessile curvedsimple

F. gothoburgensis saxicolous eastern present divaricatic acidUV+ white

grey areoles�dispersed, smooth

I� sessile straight

F. intercincta saxicolous western absent divaricatic acidUV+ white

grey, areoles smooth I�(rarely+violet)

innate straight

F. kochiana saxicolous Europe absent divaricatic acidUV+ white

grey, areoles smooth I� innate straight

F. lightfootii corticolous Europe present divaricatic acidUV+ white

greyish to green I� sessile constrictedat centre

F. lowensis saxicolous eastern absent divaricatic acidUV+ white

brown, thick,areoles convex

I+ violet sessile straight

F. mollis saxicolous western absent divaricatic acidUV+ white

grey, areoles smooth I� sessile broad-ellipsoid/subglobose

F. praeruptorum saxicolous/corticolous

eastern present alectorialic acidC & KC+ red;Pd+ yellow UV+ orange

brown or grey I� sessile bent

F. pusilla corticolous eastern andwestern

present divaricatic acidUV+ white

grey I� unknown unknown

F. recensa var.arcuatula

saxicolous/corticolous

eastern absent divaricatic acidUV+ white

brown (or grey) I� sessile curved0–1-septate

F. recensa var.recensa

saxicolous/corticolous

central present divaricatic acidUV+ white

grey (or brown),areoles convex

I� sessile curved

F. scrupulosa saxicolous eastern absent alectorialic acidC & KC+ red;Pd+ yellow UV+ orange

brown, areoles smooth I� sessile straight

F. texana saxicolous southern absent norstictic acidK+ red, Pd+ yellow

pinkish-grey, areolate I� sessile broad-ellipsoid

F. thomsonii saxicolous western absent divaricatic acidUV+ white

grey, areoles smooth I+ violet innate broad-ellipsoid

2008F

uscideain

North

Am

erica—F

ryday323

of radiating hyphae, with golden-brown,K+ brown, pigment (superba-brown). Ascicylindrical, 60–70�10–12 �m, 8-spored,hypnorum-type, similar to Porpidia-type butwith a tholus with a solid I+ blue ‘plug’,rather than a hollow ‘tube’; ascospores simple,hyaline but old spores becoming brown,�uniseriate in ascus, thick-walled, 14–15�8–9 �m.

Conidia not observed.

Chemistry. K�, C�, KC�, Pd�, UV+pale yellow. No substances by TLC (fideBrako).

Distribution and ecology. Known only fromthe type collection from a mossy, dead treetrunk (Quercus) in Florida.

Additional specimen examined. USA: Florida: [Semi-nole Co.], Sanford, on dead trunk of Quercus virginiana,3 1919, S. Rapp (F—topotype).

Key to species of Fuscidea reported from North America

(Names in bold are accepted North American taxa.)

1 Thallus with soredia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2Thallus lacking soredia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

2(1) Soredia Pd+ red, C�, KC�, UV� (fumarprotocetraric acid) or Pd+ yellow,C+ red, KC+ red, UV+ orange (alectorialic acid) . . . . . . . . . . . . . . 3

All spot tests negative, UV+ white (divaricatic acid) . . . . . . . . . . . . . . 4

3(2) Soredia Pd+ red, UV�, C�, KC� (fumarprotocetraric acid); corticolous . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. arboricola

Soredia Pd+ yellow, UV+ orange, C+ red, KC+ red, (alectorialic acid), saxi-colous or corticolous . . . . . . . . . . . . . . . . . . . . . F. praeruptorum

4(2) Corticolous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5Saxicolous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

5(4) Thallus indistinct, rarely more than 10 mm diam., greyish green to green,composed of scattered to confluent granules 0·08–0·16 mm diam; soraliaefforescent and irregular; prothallus brown, typically broad & conspicuous.Apothecia and pycnidia unknown . . . . . . . . . . . . . . . . . F. pusilla

Thallus distinct, widespreading, greyish-green to brown, composed of corticategranules 0·1–0·25 mm diam.; soralia usually well developed and ulcerous . . 6

6(5) Thallus olivaceous to dull green; soralia pale or yellowish-green; apothecia and/orpycnidia usually present . . . . . . . . . . . . . . . . . . . . . . F. lightfootii

Thallus brown to grey; soralia creamish-green, surface soralia brown; apotheciaand/or pycnidia usually absent . . . . . . . . . . . F. recensa var recensa

7(4) Thallus grey, thin, often of dispersed areoles; apothecia, when present small, withwhite ring around inner margin of proper exciple . . . . F. gothoburgensis

Thallus thicker, of contiguous areoles; apothecia, when present, without white ringaround inner margin of proper exciple . . . . . . . . . . . . . . . . . . . . . 8

8(7) Ascospores simple, globose to sub-globose, 8–9 �m long; apothecia pale brown,mostly adnate with irregular, often striate margin; disc often pruinose . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. appalachensisAscospores 0–1 septate, ellipsoid, 10–12 �m long, often curved; apothecia dark

brown to black, sessile, apothecial margin at most flexuose; disc rarely pruinose. . . . . . . . . . . . . . . . . . . . . . . . . . . . F. recensa var. recensa


9(1) Medulla with positive spot test reactions: K+ red (norstictic acid), Pd+ red(fumarprotocetraric acid), or C & KC+ red, Pd+ yellow, UV+ orange (alecto-rialic acid) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

All spot tests negative, medulla UV+ white (divaricatic acid) . . . . . . . . . 13

10(9) Medulla K+ red (acicular crystals in section), Pd+ yellow (norstictic acid). Knownfrom a single locality in southern Texas (Big Bend NP) . . . . . . F. texana

Medulla K�, Pd+ red (fumarprotocetraric acid) or Pd+ yellow (alectorialic acid). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

11(10) Medulla Pd+ red, K, C, KC & UV� (fumarprotocetraric acid); ascosporesbean-shaped. Not correctly reported from North America . . . F. cyathoides

Medulla Pd+ yellow, K�, C & KC+ red; UV+ orange (alectorialic acid);ascospores globose to sub-globose . . . . . . . . . . . . . . . . . . . . . . 12

12(11) Areoles flat. Known only from boreal north-eastern North America . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. scrupulosa

Areoles convex. Known only from Japan and the Bering Straits . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. circumflexa

13(9) Medulla I+ violet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14Medulla I� . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

14(13) Thallus thick, with convex areoles; apothecia with constricted base and cupularexciple. Mountain rocks in north-eastern North America . . . . . F. lowiana

Thallus brown or grey; apothecia innate to sessile but not constricted at base andwith annular exciple. Maritime rocks in north-western North America . . 15

15(14) Thallus grey to pale brown; apothecia �innate with poorly developed margin anda white ring around inside of margin; epihymenium and exciple pale brown .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. thomsoniiThallus pale to mid-brown; apothecia �sessile with a well developed margin and

lacking a white ring around inside of exciple; epihymenium and exciple darkbrown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. aleutica

16(13) Ascospores often curved, 10–12 �m long, often 1-septate . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . F. recensa var. arcuatula

Ascospores not curved, globose to sub-globose or broadly ellipsoid, 8·0–11·5(–12·5) �m, always simple . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

17(16) Apothecia sessile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18Apothecia innate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

18(17) Thallus verrucose. Known only from eastern North America . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. appalachensis

Thallus smooth. Known only from north-western North America . . F. mollis

19(17) Apothecia with white ring around inner margin of proper exciple . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F. intercincta

Apothecia without white ring around inner margin of proper exciple . . . . . 20

20(19) Apothecia completely innate, proper margin very thin; thallus composed of smoothareoles. Not correctly reported from North America . . . . . . . F. kochiana

Apothecia adnate with thick, often striate proper margin; thallus composed ofconvex areoles . . . . . . . . . . . . . . . . . . . . . . . . F. appalachensis


I thank the curators of ABL, ASU, BG, CANL, DUKE,E, F, FH, H, MICH, MIN, NY, QFA, S, PH,US, UPS, W and WIS for the loan of collections intheir care, and to D. Flenniken (Wooster, OH), J,Gagnon (Quebec), E. Lay (Boston), J. Lendemer(Philadelphia), G. Perlmutter (Chapel Hill, NC), andG. Thor (Uppsala), for loans from their personalherbaria. I also thank T. Tønsberg (BG) for infor-mation on his additional North American collections ofF. arboricola and F. pusilla, J. Lendemer (NY and PH)for entering label data of additional collections of F.arboricola in NY into their ‘Virtual Herbarium’ <http://sciweb.nybg.org/science2/VirtualHerbarium.asp>, andfor searching Eckfeldt’s herbarium (PH) for relevantmaterial, R. Harris (NY) for details of specimens in S.Rapp’s herbarium in FLAS, H. Deguchi (HIRO) forhelp in trying to obtain collections from Dr. Inoue, A.Aptroot (ABL) for the gift of his saxicolous Fuscideacollection from Brazil, B. Coppins (E) for correcting myLatin diagnoses, A. Dibble for the photograph used inFig. 3, and the management committee of Baxter StatePark, Maine for granting a permit to collect lichens onKatahdin.

R

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Accepted for publication 8 April 2008


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